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'."^THE JOURNAL
@NJ THE «
@F RsESlA.^CHJ
m.
Established in 1 962
Edited by WILLIAM HOVANITZ
Volume 1
1962 - 1963
f
Published at
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
THE JOURNAL OF RESEARCH
ON THE LEPIDOPTERA
is sponsored by
THE LEPIDOPTERA FOUNDATION
CONTENTS
Volume 1
Number 1
August, 1962
Editorial
1
Variation in the silvering of Argynnis (Speyena) cdlippe in
the interior mountain area of south central California . . .
Oscar Elton Sette 3
The effect of various food plants on survival and growth rate of
Pieris .... William Hovanitz and Vincent C. S. Chang 21
General characteristics of the movements of Vanessa cardui (L.)
J. W. Tffden 43
Three factors affecting larval choice of food plant
William Hovanitz and Vincent C. S. Chang 51
The generic, specific and lower category names of the Nearctic
butterflies. Part 1 •— The genus Pieris . . Paddy McHenry 63
The distribution of the species of the genus Pieris in North
America William Hovanitz 73
Did the caterpillar exterminate the giant reptile? ....
S. E. Flanders 85
Further evidence of the distribution of some boreal Lepidoptera
in the Sierra Nevada Clyde Eriksen 89
Argynnis and Speyer ia . William Hovanitz 94
Volume 1 Number 2 January, 1963
Composition and relative abundance in a temperate zone
butterfly fauna . . . Thomas C. Emmel and John F. Emmel 97
The Argynnis populations of the Sand Creek area, Klamath
Co., Oregon, Part I .... J. W. Tilden 109
Caterpillar Versus Dinosaur? . . . Theodore H. Eaton, Jr. 114
Geographical distribution and variation of the Genus Argynnis
I. Introduction
II. Argynnis idalia William Hovanitz 117
The relation of Pieris virginiensis Edw. to Pieris napi L.
Species formation in Pieris? ..... Wiliam Hovanitz 124
The male genitalia of some Co lias species . Bjorn Petersen 135
The effect of hybridization of host-plant strains on growth rate
and mortality of Pieris rapae .
WiUiam Hovanitz and Vincent C. S. Chang 157
Notice-editorial
162
Change of food plant preference by larvae of Pieris rapae
controlled by strain selection, and the inheritance of this trait
William Hovanitz and Vincent C. S. Chang 163
Volume 1 Number 3 March, 1963
Selection of allyl isothiocyanate by larvae of Pieris rapae and
the inheritance of this trait
William Hovanitz and Vincent C. S. Chang 169
Biology of the Ceanothus stem-gall moth, Periploca ceanothiella
(Cosens), with consideration of its control. J. Alex Munro 183
Larval food-plant records for six western Papilios, ....
John F. Emmel and ThtMnas C. Emmel 191
CoUas philodice in Chiapa^ Mexico . . Thomas C. Emmel 194
Notes on the early stages of two California geometrids
John Adams Comstock 195
Geographical distribution and variation of the genus Argynnis
III. Argynnis diana William Hovanitz 201
The generic, specific and lower category names of the nearctic
butterflies. Part 2 — The Genus Colias . Paddy McHenry 209
A standard method for mounting whole adult lepidoptera on
slides utilizing polystyrene plastic . . Charles L. Hogue 223
Volume 1 Number A May, 1963
Techniques in the study of population structure in
Philotes sonorensis
Rudolf H. T. Mattoni and Marvin S. B. Seiger 237
Early stages of a southern California Geometrid moth,
Drepanulafrix hulsti hulsti (Dyar) John Adams Comstock 245
The effeaiveness of different isothiocyanates on attraaing larvae
of Pieris rapae
William Hovanitz, Vincent C. S. Chang and Gerald Honch 249
The origin of a sympatric species in Colias through the aid of
natural hybridization William Hovanitz 261
A method for breeding Pieris napi and Pieris bryoniae
Bjorn Petersen 275
An analysis of the North American species of the genus
Callophrys J. w. Tilden 281
THE LEPIDOPTERA FOUNDATION 301
Notice 302
Volume 1
Number 1
August, 1962
Established in 1 962
Published at
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Edited by WILLIAM HOVANITZ
WITH EMPHASIS ON ENVIRONMENTALLY AND GENE-
TICALLY INDUCED VARIATION, population analysis, evolution,
phylogenetic taxonomy, zoogeography, comparative morphology, ecol-
ogy, geographical variation, speciation, physiology, etc. In short,
quality tuork on any aspect of research on the Lepidoptera.
THE PURPOSE OE THE JOURNAL is to combine in one source
the work in this field for the aid of students of this group of insects
in a way not at present available. The JOURNAL will attempt to
publish primarily only critical and complete papers of an analytical
nature, though there will be a limited section devoted to shorter
papers and notes.
RATES: $ 8.00 per volume, personal subscription.
$12.00 per volume, institutional subscription.
Benefactor subscriptions are encouraged. These may be
any amount over those above.
All amounts are in U.S. dollars, payable in the U.S.A.
AUTHORS ARE REQUESTED to refer to the Journal as an
example of the form to be used in preparing their manuscripts. Fifty
separates will be supplied to authors free; reprints will be sold at
printer’s rates if ordered at time galley proofs are returned. If proofs
are not returned promptly, the editor reserves the right to withhold
publication, or to proceed with no responsibility. All issues of the
Journal will be copyrighted; in submitting a paper each author agrees
that the material has not been published elsewhere.
MANUSCRIPTS AND SUBSCRIPTIONS should be mailed to the
address noted above.
BENEFACTORS Individuals or institutions who by reason of
their interest in the welfare or research on the Lepidoptera, and in
its publication, who wish to contribute beyond the cost of a subscription,
will be listed here.
BIO-METAL ASSOCIATES
THOMAS C. EMMEL
ANONYMOUS DONOR
ANONYMOUS DONOR
1(1):1,1962
Journal of Research on the Lepidoptera
1140 W. Orange Grove Ave., Arcadia, California, U,S.A.
Copyright 1962
EDITORIAL
Starting a new journal is not an easy matter, nor is it a matter free
of controversy. This is especially the case where a new journal, as this
one, is started without the support of an existing society. This journal
is intended as a meeting ground for papers on all aspects of the biology
of the Lepidoptera, moths and butterflies alike. Too many entomolog-
ical journals don’t offer publication of articles of the length necessary
to present sound data and Completeness of effort. The Lepidoptera are
a group of insects second to none in the scope of our knowledge about
them: work in this fieM has reached the higher levels of learning and a
journal is required to keep pace with this knowledge at the same level.
A journal with a wide scope is our aim — narrow in the taxonomic
sense but wide in the biological sense. The journal will attempt to
publish primarily only critical and complete papers of an analytical na-
ture, though there will be a limited section devoted to shorter papers
and notes. There will be emphasis on environmentally and genetically
induced variation, population analysis, evolution, phylogenetic taxon-
omy, zoo-geography, comparative morphology, ecology, geographical
variation, speciation, physiology, etc.
It has been very gratifying to receive so many subscriptions on the
basis of merely a brochure; every effort will be made to keep the journal
up to the standard expected of us. The response in so short a time has
served to show those who have generously provided the finances that a
journal of this type has been needed and will fill a desirable niche.
We are happy to publish in this first issue a number of papers
which the authors were able to get ready on "short notice". There is
a preponderance on one taxonomic group, but with different points in
mind. With this first issue out, we offer the pages of this journal to all
with papers to publish on the Lepidoptera that fit the intent of the
journal. A wider range of papers will come in future issues.
1
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l(l):3-20, 1962
Journal of Research on the Lepidoptera
1140 W. Orange Grove Ave,, Arcadia, California, U.S.A.
Copyright 1962
VARIATION IN THE SILVERING OF ARGYNNIS
(SPEYERIA) CALLIPPE IN THE INTERIOR
MOUNTAIN AREA OF
SOUTH CENTRAL CALIFORNIA
OSCAR ELTON SETTE
23643 Arbor Avenue, Los Altos, California
In a comprehensive study of the geographic variation of
Argynnis (Speyeria) callippe Boisduval, based on examination of
about 2750 specimens from many localities well distributed through
out the range of the species in California, Hovanitz (1943) recog-
nized several main divisions of this species complex. Among them
are the *'South Coast Range Population” extending from the San Fran-
cisco Bay area southward through the Coast Ranges into Lower Calif-
ornia and the "Western Sierra Nevada Populations” extending in a
band along the western Sierra Nevada, at moderate elevations from
the region west of Lassen Peak southward to about the Greenhorn
Mountains at the southern end of the Sierra Nevada. He found that
these two population groups are connected through a "Southern Zone
of Intergradation” by a "series of steps across the Piute Mountains,
the Tehachapi Mountains and the Sierra Madre Range.”’
He characterizes the populations in the Southern Zone of Inter-
gradation as follows:
"From the Santa Monica Mountains ‘ on the coast, where the typical
callippe of this (the South Coast Range) region lives, it is found that
in going inland (Charlton Flat, Mint Canyon and ’Ridge Route’) the
lightness of all colors increases. The butterfly becomes smaller and the
light colored band and spots on the upper surface of the wings tend to
become obliterated, leaving a more uniformly colored wing surface such
as is present in the western Sierra Nevada gradient. However, the yellow-
brown color is very much lighter than in the latter and the band on
the under side of the hind wings is still yellow; the spots are still always
fully silvered. The tendency toward these conditions is the more marked
the farther from the coast and the farther into the Tehachapi Range
the populations exist. In the Tehachapi Range, the butterflies are very
lightly colored and the band on the upper surface of the wings is rare;
the spots are still silvered. At Havilah, Piute Mountains, the population
consists of some silvered, some unsilvered and some intermediate spotted
IThis range is not named on many maps. It lies parallel to, and south of, the middle portion
of the Cuyama River which is the long north branch of the shorter Santa Mafia River shown
discharging into the ocean at about lat 35° in Fig. 1.
3
4
SETTE
J. Res. Lepid,
individuals (this is the type locality of macaria Edws.); the exact fre-
quency of these types is not known; but there is a high percentage of
silvered and unsilvered present ... In the Greenhorn Mountains, the
segregation into a silvered population on the eastern side of the summit
and an unsilvered one on the western side is decidedly apparent, though
mixing occurs toward the south, where the populations unite”.
The main purpose of this paper is to describe semi-quantitatively
the variation of silvering within the Southern Zone of Intergradation.
The silvery decoration of animals is far less common than the pigmen-
tal coloration. While Hovanitz ( 1941 ) has found general qualitative
correlations of the pigmental coloration of ^butterflies with environ-
mental conditions, he finds the significance of silvering in callippe
"quite incomprehensible" (Hovanitz, 1943, p. 420). Some peculiari-
ties of the silvering data revealed by the semi -quantitative treatment
in this paper suggests that studies of its physico-chemical nature and
genetic control might be rewarding. An hypothesis is advanced as a
point of departure for such studies.
THE SAMPLES
The material for this study consists of complete series of callippe
that I collected during June 8 to 15, 1957 in company with Fred T.
Thorne, and one sample from Bouquet Canyon consisting of part of
a series taken in 1952 by Thorne and kindly given to me. In brok-
ing the series Thorne exercised no conscious selection for color or
pattern, though worn and damaged specimens were discarded.
Parallel or convergent evolution among the Argynnids in some
regions presents problems of species identification. In the Southern
Zone of Intergradation this problem arises between calliiype and cor-
onis Behr. The latter, as the subspecies hennei Gunder, flies with cal-
lippe and resembles it so closely that confident identification requires
either great familiarity with both species or known series from the
region for comparison. L. P. Grey is in possession of both and kindly
reviewed my field identifications. These had been made in consulta-
tion with Thorne. At the time neither of us had previous acquain-
tance with hennei, though we were both familiar with several other
subspecies of coronis. Two specimens I had labelled callippe proved
to be undoubted coronis. One labelled callippe came back from Grey
with the notation, "Could this be coronis? (Don’t ask me!." Another
labelled coronis bore Grey’s notation "I think it is a callippe!* The
last two, when spread and compared with the now-available good
series of hennei, appear to me to be undoubted coronis, though the
under side wing surface (the only surface exposed to Grey’s examina-
tion) is remarkably like the callippe from the same locality. These
four speciments have been excluded from the samples.
In the list that follows, the collections are grouped by localities.
Each group contains mainly specimens taken at one location, to
which have been added a few specimens from nearby locations. Those
i(j):}-20, 1^62
VARIATION IN SILVERING
5
in the main group, I believe, are members of a single interbreeding
colony; the others may consist of strays from the same colony in some
instances, and in others, may be strays from some other colony. All
localities are in Kern County except Sandberg’s and Bouquet Canyon
which are in Los Angeles County. Distances from named places are
given to the nearest mile (I.6I km) measured on a straight line on
large-scale road maps. Directions, also determined from such maps,
are referenced to true North and stared in Mariners’ abbreviations
appropriate to the 32 -point compass. Thus southwest is siven as SW,
southwest by west as SWxW and west southwest as WSW. These are
11.25° steps and allowing for measuring inaccuracy are correct to
about the nearest 15°. Elevations were taken by aneroid altimeter
graduated in 100-foot (30.5 m) units and probably accurate to ±
200 feet. Botanical names are according to Jepson (1923 - 1925),
Abrams (1940, 1944 and 1951) and Abrams and Ferris (I960). No-
menclature of the butterflies follows the list by McDunnough (1938)
except for the genus {Speyeria), which, at the species and subspecies
level follows the arrangement of Dos Passos and Grey (1947).
A. East slope, Greenhorn Mountains. June 12, 1957, 18 males and
1 female taken 6 miles WSW of Kernville, elevation 4900 ft., along the road
leading steeply from Isabella Reservoir to Greenhorn Mountain Park passing
between Tittle and Rattlesnake Creeks, in association of Digger Pine (Pinus
sabiniana Dough), Oak (Quercus sp), California Fremontia (Premontia cali-
fornica Torr.), and chaparral broken by grassy areas, with Yerba Santa {Brio-
dictyon so.) flowers as nectar attractant; included in addition is 1 male taken
8 miles WxS of Kernville, elevation 5500 ft., Transition Zone, in a small
roadside opening in Yellow Pine (Pinus ponder osa Dough) and Incense
Cedar {Libocedrus decurrens Torr.) forest.
B. West slope, Greenhorn Mountains. June 12, 1957, 58 males and
6 females taken 27 miles NE of Bakersfield, elevation 3300 ft., near the foot
of Eugene Grade, in Digger Pine, Blue Oak (Qurcus douglasii Hook & Arn.)
and grass association, with violets (Viola sp.) abundant; included in addition
are 3 males and 1 female taken 26 miles NNE of Bakersfield, elevation 3000
ft., in Blue Oak and grass association, with riparian flora along a small creek.
C. Havilah. June 11, 1957, 13 males and 1 female taken 2 miles S of
Havilah, elevation 3100 ft., Digger Pine, mixed oak and chaparral association
with riparian flora along a small creek, mostly attracted to Yerb Santa flowers;
in addition are 1 male from 1 mile north of Havilah, elevation 2800 ft., and
another from 5 miles S of Havilah, elevation 3900 ft.
D. Walker Basin. June 11, 1957, 30 males and 6 females taken 10 miles
SxW of Havilah, at the south end of Walker Basin, elevation 3200 ft.. Blue
Ok and Grass association with sagebrush and violets. Hoarhound (Marrubium
vulgare L.) attracted a few individuals, but most were taken in the characteristic
slow, fluttering flight displayed when on breeding grounds.
E. Tehachapi. June 14, 1957, 12 males and 7 females taken 8 miles W
of Tehachapi (the town), elevation 4700 ft, at the eastern end of Bear Valley
in Blue Oak and grass association, with patches of sagebrush (Artemisia tri-
dentata Nutt.), mostly acctracted to Wallflower (Erysimum sp.) flowers; in
addition are 3 males from 7 miles W of Tehachapi, elevation 4600 ft., and 1
female from 3 miles SW of Tehachapi, elevation 4600 ft.
6
SETTE
/. Res. tepid.
I2 0‘
119*
118®
36'
35*
3 4*
FIGURE 1. Location of populations sampled (letters "A to H”) super-
imposed on the distribution of Argynnis (Speyeria) callippe (shaded area)
adapted from a portion of Hovanitz’ map (1943, Fig. 1) showing the distribu-
tion of the "silvered spots” character by shading lines slanted upward to the
left and the "unsilvered spots” character by shading lines slanted upward to
the right. The Kern River (upper right) separates the Greenhorn Mountains
to the north from the Piute Mountains to the south as it flows to a sink in the
lower San Joaquin Valley (blank area at upper middle). At middle left is the
Santa Maria River with the Cuyama River as its long north fork. At the lower
right is the Santa Clara River. The crests of the higher mountains and ridges
of the transverse series of ranges lie north of the Santa Clara River Drainage and
south of the Cuyama River.
F. Lebec, June 8, 1957, 24 males and 8 females; June 9, 1957, 26 males and
4 females; and June 15, 1957, 11 males and 15 males; total, 61 males and
27 females taken 1 mile S of Lebec, elevation 3600 ft., in oak and grass
association with adjacent chaparral including sagebrush {A. tridentata) , Buck-
wheat Brush {Eriogonum^ fasciculatum Benth.) and Yerba Santa; some were
at flowers of the last, others in breeding-ground flight.
(r;.-j-20, 1^62
VARIATION IN SILVERING
7
G. Sandberg^S (on old Ridge Route). June 9, 1957, 12 males and 3 females;
June 15, 1957, 1 male and 2 females; total, 13 males and 5 females taken 9
miles ESE of Gorman in Digger Pine and sagebrush (not A. tridentata) associa-
tion, with violets. Breeding ground behavior was predominant.
H. Bouquet Canyon. June 11 and 12, 1951, 18 males collected by Fred
T. Thorne at "upper Bouquet Canyon, 4 miles down from Pine Creek — Lake
Hughes road” (estimated 16 miles W of Palmdale, estimated elevation
3400 ft.).
HABITAT AND MESOSCALE DISTRIBUTION
According to these records callippe in this region occupies ele-
vations from 2800_ to 5500 feet, but occurs in good concentrations only
between 3100 and 4900 feet, generally, the upper portion of the Up-
per Sonoran Life Zone. Its habitat is marked characteristically by scat-
tered Blue Oak, Digger Pine, and sparse chaparral with sagebrush {A.
tridentata) often pesent. Usually there are broad areas or small patches
of grass present. Practically all of the localities are in grazing land,
but at the time of our trip it was being very lightly grazed.
Violets were noted at only three of the locations. Where they were
seen, the plants were abundant, mature, with fully-developed seed cap-
sules and appeared about to dry up. Where violets were not seen,
either they had dried up already, or the collecting was of members of
the colony that had been drawn by nectar sources somewhat away
from the site of the larval food plant.
During the trip we collected at 31 locations within the geographic
range and elevation limits of this butterfly. Of these, seven obviously
were in, or close to, callip'be colonies; five locations yielding only one
or several specimens, evidently were not as close; and the remaining
19 locations where we took none probably were distant from colonies,
though this may not have been true for a few places where we sighted
an Argynnid or two v/hich could have been either this species or coro-
nis. These indications favor a population model consisting of well-
separated, compact colonies from which individuals do not stray often
or far. This pattern is more accentuated in this region than in most
others within my collecting experience.
Butterflies of 30 other species were taken at one or another of the
seven locations where callippe were caught in good numbers. Plebeius
acmon West, and Hew. and Hesperia lindseyi Holland were the most
ubiquitous, occuring at most locations from the Greenhorn Mountains
to Sandbergs. The former were taken in small numbers while the latr
ter, in some locations were very abundant. In addition to these, the
most common and prevalent in the Greenhorns and the Piutes were
Euphydryas chalcedona Doubleday and Hewitt, Melitaea palla Boisdu-
val, and Strymon saepl^m^ Boisduval; while through the Tehachapi to
Sandberg’s they were Minois silvestris Edwards and Argynnis coronis
Behr. If one of the specimens whose identification was questioned by
8
SETTE
/. Res. Lepid.
Grey is, as I believe, coronis, this species extended north at least to
Havilah. Inasmuch as all coronis taken on this trip were fresh males,
and in other regions within my collecting experience coronis flies some-
what later than callippe, it is likely that a week or two later in the
season coronis would have been found to be even more consistently
a companion of callippe.
RELATIVE ABUNDANCE
The relative abundance in butterfly populations may be signifi-
cant in studies of variation. From records extending through many
years, Ford (1945) found that a rapid increase in abundance of the
colonial butterflv Euphydryas aurina Rott was accompanied by "an ex-
traordinary outburst of variation". When the increase ceased, the
colony settled down to a comparatively uniform type, different from
the one prevailing prior to the increase. Scientists differ as to the in-
terpretation and significance of this phenomenon as a mechanism in
evolution. Whatever the interpretation, if the generality of this phen-
omenon is to be ascertained, records must be accumulated through the
years for a number of butterfly populations.
To this end, there are placed on record in Table 1, estimates of
relative abundance of callippe in each of the seven colonies sampled
in 1957. This estimate is in terms of catching rate and assumes that
the number caught in unit time is proportional to the abundance of
the population in the area of collecting. It is homologous in concept
to the measure "catch per unit of effort” almost universally and suc-
cessfully (sometimes "calibrated” to a absolute abundance by tag-and-
recapture experiments) used as a basic statistic in studying the dy-
namics of fish population fluctuations (Ricker 1940). For callippe
I have computed the catching rate, R, according to the formula:
R ^ Nc .
H - k Nt
where Nc is the number of callippe caught, H is the time, in hours,
in the collecting period, Nt is the total number of specimens of all
species caught (including callippe) during the collecting period, and
k is a constant whose value depends on the methods and dexterity of
the collector in caring for a specimen once it is in the net. By timing
myself in the_caring, individually, for each of 150 specimens of a
number of species from all families except Megathymidae under typ-
ical field conditions, I found my k value to be 0.0155 hours (37.8
seconds ) .
In repeated samplings of several species I have fo*und R to be sur-
prisingly stable. For callippe, the three samplings at Lebec are an ex-
ample of this. Although stable, R as computed by this formula should
i(i):3’2o, I $62
VARIATION IN SILVERING
9
be regarded only as a first-order approximation, or index, of true rel-
ative abundance, because it is affected by systematic error and several
sources of variability, not all of them random. A more sophisticated
iterative treatment theoretically should improve R, but it involves as-
sumptions of uncertain merit awaiting test.
TABLE 1. Relative abundance in terms of catching rate, R, in number per
males and by the per cent of the males that were fresh.
Catch-
ing Per Per cent
Locality, date (1957) and elevation rate cent of males
(R) males fresh
A. E. slope, Greenhorns, el. 4900 ft., June 12 ....... * 39 84 86
B. W. slope, Greenhorns, el. 330 ft., June 12 ........ **227 91 52
C. Havilah, el. 3100ft., June 11 16 93 15
D. Walker Basin, el. 3200 ft., June 11 * 44 83 87
E. Tehachapi, el. 4600 - 4700 ft., June 14 33 67 79
F. Lebec, el. 3600 ft., June 8 24 75 84
F. Lebec, el. 3600 ft., June 9 24 87 69
F. Lebec, el. 3600 ft., June 15 30 42 45
G. Sandberg’s, el, 4000 ft., June 9 17 80 75
In the meantime, of most concern is that R increasingly underes-
timates abundance as the latter increases, presenting simultaneous
catching opportunities more often and usually avail can be taken of
only one of them at a time. Recognizing this, the values in Table 1
that probably were moderately underestimated owing to this factor
are marked with an asterisk, and the one value that was grossly under-
estimated is marked with a double asterisk. At the other end of the
scale, when only one or two specimens are caught during a collecting
period, R has large error owing to random variability of incidences
of encounters. Avoiding this. Table 1 gives catching rates computed
for only the first-listed location under each locality, except for Tehach-
api where the times and catches of the first two locations were pooled
for computing the catching rate.
Another error source is that as kNt approaches H, a small error
in k produces a large error in R. This approach was not close enough
to be critical for the values given in Table 1, though the value for the
west slope of the Greenhorn Mountains may have been moderately
affected.
Every collector can think of a number of other obvious sources of
error or variability such as weather conditions, unusual concentrations
at attractants, etc.. These obvious ones can be avoided by comparing
only sets of samplings taken under reasonably similar conditions, as
is true for the set in Table 1 except as later noted.
Of course the catching rate reflects the relative abundance only at
the stage of flight at which the collecting was done, and may yield
values departing substantially from the inherent abundance. An esti-
10
SETTE
/. Re$. tepid.
mate of the stage of the flight is afforded by the relative numbers of
fresh and worn individuals. In Table 1, recorded as *'fresh” were in-
dividuals without readily perceptible random loss of scales or fringe.
Damage reasonably attributable to accidental causes, such as notches,
tears, breaks and rubbed streaks or patches was not considered. The
data are given for males only, there not being enough females to yield
reliable percentages. The two sexes were not combined for this sta-
tistic because this would introduce extraneous variability owing to the
tendency of females to emerge later than males. As evidence of this,
only four of the 53 females listed in Table 2 were worn. More direct
evidence is afforded by the Lebec samples. Those of June 8 and 9 were
mostly males and mostly fresh. About a week later more than half of
the males were worn and the females, all fresh, outnumbered the males.
The sample from Havilah is anomalous in respect of percentage worn
in relation to sex ratio. Although most of the males were worn, only
one female was taken. Probably the greater ease of collecting males at
the Yerba Santa flowers diverted us from the females which probably
were widely scattered for ovipositing in the adjacent area which ap-
peared to be the likely habitat for violets, though the plants were not
observed, doubtless having dried up before our visit. At Sandberg’s
the catching rate was substantially depressed bv a strong breeze sweep-
ing the exposed slope and wafting many a disturbed butterfly out of
stalking or pursuit range.
Appraising the catching rates in the light of these qualifying fac-
tors, the presence of some worn males at all localities indicates that the
callippe flight was well developed throughout the region. But the high
percentage of fresh males and the scarcity of females suggests that the
flight had not yet reached its peak at most localities except Lebec where
it probably was near peak stage on June 15 and Havilah, where the
male flight had apparently passed its peak. Allowing for the differ-
ences in stage of flight and the collecting difficulties at Sandberg’s, it
appears that relative abundance was capable of attaining a height-of-
flight index of 30 to 40, probably closer to the latter, at all localities
except the location on the west slope of the Greenhorn Mountains
where abundance was at least an order of magnitude higher, and at
Walker Basin where it probably was substantially higher than 40 per
hour, though well below that on the west slope of the Greenhorn Moun-
tains. Although we did not trace out the geographic extent of a colony
in any of the localities, the impression gained while collecting, was
that these two colonies covered more extensive areas than the rest. It
may be concluded, first, that the samples were drawn from near the
middle of the flight period and hence probably represent nearlv the
modal characteristics of the population in each colony; and second, the
colonies varied quite widely in population size, perhaps through more
than one order of magnitude. Further results may emerge if compar-
isons with changed levels of abundance become possible in the future.
1^62
VARIATION IN SILVERING
11
VARIATION IN SILVER SCALING
The silver scaling in callippe^ as in Argynnids generally, is confiined
to certain pattern elements on the ventral wing surface. For the most
part these are well defined spots in the apical and subapical area of
the fore wing and all well defined spots in all areas of the hind wing.
In addition, on heavily silvered individuals, silver scales may form ill
defined streaks along the inner margin and in some of the interspaces
between spots in the basal and discal areas of the hind wing. The pres-
ent study is confined to the well defined spots of the hind wing. When
unsilvered these spots are a light buff color, usually sharply bordered
or outlined by brown scales. Some or all of the spots may be partly sil-
vered. Then the silver scales and buff scales stand out in sharp con-
trast to each other when specimens are held in the best relation to the
light source and the eye to bring out the specular quality of the silver
scales.
A fully quantitative measure of the degree of silvering would be
the ratio of silvered scales to all scales in the area potentially subject
to silvering. To avoid spending a prohibitive amount of time count-
ing scales, I have employed a subjective system of scoring based on
general appearance. It would have been desirable, too, to utilize ex-
clusively the pristine fresh individuals, but to avoid reducing the num-
ber below levels needed for tests of significance, I have scored the
slightly and moderately worn individuals along with the fresh ones, ex-
cluding only those whose loss of scales caused serious doubt as to their
correct score. The penultimate column of Table 2 shows the number
excluded for this reason.
Because the amount' of silvering sometimes differed appreciably in
the basal and discal area, which will be called "disc” for brevity, from
that in the submarginal area, called "margin”, the two portions of the
wing were scored separately. Four grades of silvering were recognized:
Grade 0 for complete lack of silver scales: Grade 1, when there were
only flecks of silver well separated by buff scales; Grade 2, when some
of the spots were unsilvered, others were partly silvered and still others
were fully silvered, or any combination of two of these conditions; and
Grade 3, when all spots were so fully silvered that there were no readily
perceptible buff scales. The Disc Grade and the Margin Grade were
treated as half scores and added together to arrive at the individual
Score. Thus there are seven possible scores: 0 to 6, inclusive.
Of the 238 individuals scored, 50 had half scores that differed from
each other by one grade point. None differed by more than one. Of
the 50 with different half scores, the Disc Grade was higher than the
Margin Grade for 40 individuals and lower for 10. Curiously, for the
two Greenhorn Mountain samples and the' Havilah sample, pooled, the
ratio was 33:4, while for the Walker Basin, Tehachapl and Lebec sam-
ples, also pooled, it was 7:6. The difference between the ratios 33:4
12
SETTE
/, Re$. tepid.
and 7:6 is statistically significant (p=0 02) desoite the small numbers
in the latter. It is possible that the tendency for the marginal row of
spots to be less silvered than the disc spots is linked to the general lack
of silvering in the three northernmost localities.
TABLE 2. Frequency distribution of silvering scores
Sex and sample Score
0 1 ' 2 3 4 5 6 Mean Scored Total
Males :
A. E. slope. Greenhorns 5 3 1 1—2 1 3.16 — - 19
B. W. slope, Greenhorns 17 8 7 64 8 7 2.42 7 64
C. Havilah 2 1 — 2 — 2 4 3.73 4 15
D. Walker Basin 1 2 1 — 1 2 19 5.08 4 30
E. Tehachapi 1 — — - i— _ — 13 5.40 — 15
F. Lebec 1 1 1 1 1 1 39 5.54 15 60
G. Sandberg’s — — — — — . — 10 6.00 3 13
Total 27 15 10 11 6 15 99 4.16 33 216
Females :
A. E. slope, Greenhorns — 1- — ■ — — — -2.00 — ■ 1
B. W. slope. Greenhorns — — 2 3 - — — — 2.60 5
C. Havilah l l
D. Walker Basin _ „ _ 11 1 3 5.00 — 6
E. Tehachapi _ _ l 1 6 5.01 — 8
F. Lebec 1 _ _ _ 25 5.77 1 27
G. Sandberg’s - — — — . — — — 3 6.00 — 5
Total 1 — 4 5 1 i 39 5.12 2 53
The mean score for males is 4.16 and for females is 5.12 (Table 2),
suggesting that females tend toward more silvering than males. But
four-fifths of the females were taken at Walker Basin, Tehachapi and
Lebec where silvering in both sexes is nearly complete. If we pool
these three sampls and cHss together the individuals with scores 0 to
4 inclusive, and similarly those with scores 5 and 6, in order to have
high enough numbers for statistical test, we find no significant dif-
ference between the sexes (P = 0.9). Accordingly, further analysis
deals with both sexes combined, as shown graphicallv in Fig. 2.
The arrangement of samples in Fig. 2 is in ascending order of their
means. This also arranges them from north to south except for the
reversal of samples A and B. Where samples are near the same par-
allel of Latitude and have appreciable meridianal displacement, as
among A, B and C as one group and F, G, and H as another, their
means also ascend from west to east.
The slopes of the lines connecting the means in Fig. 2 should not
be taken as representing gradients of silvering in the sense of reflecting
unit increase of silvering per unit distance. The panels are not spaced
in proportion to distance, but merelv to accommodate the height of the
bars. The means, themselves, are defective because the scores do not
represent even gradations in amount of silvering. Further, because of
t§€a
VARIATION IN SILVERING
13
wesr SLO PE. GREEN -
HORN MTS. 35* 37'
EAST SLOPE, GREEN -
HORN MTS. 35® 43 '
HAVILAH 35® 31'
WALKER BASIN
3 5* 23'
TEHACHAP! 35®08'
LEBEC 34*50*
SA NDBERG'S
34® 45'
BOUQ UET CANYON
34® 35'
FIGURE 2. The degree of silvering of the several samples, shown as histo-
grams of the number of individuals according to grade (see text); and the
mean grades, shown as small circles connected with heavy straight lines.
NUMBER OF INDIVIDUALS
14
SETTE
J. Res. Lepid.
the prevalence of low numbers at the middle scores, the distributions
depart widely from the bell-shaped normal probability curve and the
par? metric test commonly applied to means would not yield reliable
probabilities even if the means were not otherwise defective. The
means were computed and connecting lines were supplied in the graph
only to give a general perspective of the amount and direction of dif-
ferences between localities.
For a more careful assessment, the significance of the differences
between samples, as to the numbers of individuals falling into the sev-
eral silvering classes, can be made by the chi -square test which re-
quires neither a normal distribution nor uniform class size. It does
require a pooling of the original score classes to provide a minimum
of five as the expected number on a class. To meet this requirement,
the data have been grouped into two classes: (1) individuals with
Scores 0 - 4, which can be described as "unsilvered” and partly silvered”;
and ( 2 ) those with Scores 5 and 6, which can be described as "silvered
or nearly so”. With this grouping the approximate probability, P. of
the difference between any two samples in respect of the ratio of "un-
silvered and partly silvered” to "silvered or nearly so” occurring by
chance can be found by entering the table of chi-square distribution
on one degree of freedom with the value of chi scfuare computed from
the 2x2 contingency table according to the method given by Fisher
(1948, p. 85). For greater precision the Yates correction for contin-
uity was incorporated and P was found by graphical interpolation be-
tween tabular values. The results are given in Table 3 as a matrix in
which the chi-square values betyeen all combinations of pairs of sam-
ples are shown in the upper right portion and the corresponding values
of P in the lower left portion.
TABLE 3. Chi-square values (upper right) and probability values (lower left)
between pairs of samples.
Locality
Sample
and sample
B
A
C
D
E
F
G
W. Greenhorns ..
B
2.37
2.97
>10
>10
>10
>10
E. Greenhorns ..
A
0.16
0.02
4.63
1.10
>10
7.14
Havilah
C
0.08
0.10
1.24
1.74
0.83
1.78
Walker Basin
D
< .001
.03
.19
.01
.84
2.32
Tehachapi
E
<.001
.02
.18
.10
.64
1.36
Lebec
F
<.001
<.001
.34
.34
.40
.37
Sandberg’s —
G
<.001
.008
.02
.14
.19
.10
The three northern samples. A, B, and C form a group within which
there is no statistically significant difference at the five per cent prob-
ability level (P = 0.05), and the four southern samples, D to G, in-
clusive, form another such group. But these groups are not discrete
from each other. Sample C from Havilah differs no more significantly
from D, E, and F than from A and B, indeed, somewhat less. The sum
of chi squares among A, B and C is 5.36; df are 3; and P is 0.14. The
i(i):3-20, 1962
VARIATION IN SILVERING
15
comparable values among C, D, E, and F are: Chi-square, 5.34; df, 6;
P, 0.48. The indeterminate position of Havilah relative to the two
groups, most probably is due to the small number in the sample from
there. With such a small sample it would require a very strong con-
trast in silvering ratio to prove statistical significance. Until more ma-
terial is available it remains uncertain whether the population at Havi-
lah resembles more closely those to the North or those to the South in
respect of silvering, or whether it is truly intermediate as suggsted by
Fig. 2.
Likewise, the sample from the east slope of the Greenhorn Moun-
tains is small and does not test significantly different from that of the
west slope. But for the Greenhorn Mountains there are two sources of
additional data. Thorne and I scored both his catch and mine while
in the Greenhorns and this record includes about twice as many indivi-
duals as given in Table 2. We used three categories: ' unsilvered”,
*'part silvered”, and “silvered”, corresponding somewhat inexactly to
the Scores 0 - 1, 2 - 4 and 5 - 6, respectively, of Table 2. Hovanitz
(1934) reported the number of individuals in a sample from the east
side of the Greenhorn Mountains, elevation 5500 feet, and from the
west side at Cedar Creek, elevation 5000 feet, according to three cate-
gories: “not silvered or very slightly so”, “intermediate” and “well -
or fairly-well silvered”. Assuming these categories correspond approx-
imately to those of Thorne-Sette, the data may be pooled as follows:
East side:
Unsilvered
Intermediate
Silvered
Total
Thorne - Sette .
......6
.15
17
38
Hovanitz
....0 ......
..2
14
16
Total
West side:
...........6
17 .........
31
...54
Thorne - Sette ...
72
...24
32
128
Hovanitz
....10
8
4
22
Total ..............
.........82 ......
..........32
36
150
With more data it is possible to use a 2 x 3 contingency table for
a more discriminating test. This yields a probability far less than one
in a thousand that the west side and east side samples could have been
drawn from a population containing the same proportions of unsil-
vered, partly, silvered, and silvered members. This does not prove
that there is no mixing between the populations of the west and east
slopes; it only indicates the extreme unlikelihood of enough inter-
change to form a thorough mixture. But when it is considered, in
addition, that the ridge of the mountain is clothed with transition
forest within which no colonies of callippe were found, it seems clear
that the interchange, if any, must be slight and probably by way of
mixing with the more silvered populations to the South at different
rates on the two sides of the mountain range as suggested by Hovan-
itz (1943, p. 411).
16
SETTE
/. Res. tepid.
Turning southward, past Havilah, there is at Walker Basin a small
proportion of unsilvered and partly silvered individuals in the pop-
ulation. The low proportion continues with only slight further dim-
inution in going from Walker Basin in the Piute Mountains, through
the Tehachapi Mountains to Lebec. This uniformity suggests a rela-
tively brisk interchange between colonies in this stretch of mountains,
probably coupled with strong environmental selection against unsil-
vered individuals.
Then at Sandberg’s, only a short distance beyond Lebec, all indiv-
iduals are silvered, This is true also for the specimens from Bouquet
Canyon (not included in the scoring table because they were only a
part of a series). But my sample from Sandberg’s is too small to
assuredly include unsilvered or partly silvered individuals if they con-
stitute only a small percentage of the population. There is historical
evidence, in Gunder’s (1930) list of butterflies of Los Angeles Coun-
ty, of the occurance of unsilvered callippe at several localities in the
transverse ranges south and southeast of Lebec. Under ”Araynnis ma-
caria lamina Wri. THE UNSILVERED MACARIA FRITILLARY, a
transition form.” Gunder recorded one male collected by Comstock
"June 10, 1922, Ridge Route;” one male collected by Friday "June 19,
1929, Pine Canyon;” and two males and one female collected by Gun-
der "June 25, 1921, Bouquet Canyon.” "Ridge Route” probably is
identical to, or at least near, the locality I give as "Sandberg’s”; "Bouquet
Canyon” may be identical to, or near the locality of the Thorne col-
lection here listed under the same place name; Pine Valley lies about half-
way between the two. The last is definitely on the desert side of the
transverse ranges, while the other two localities are near the crest divid-
ing the drainage to the desert from the drainage to the Pacific. The
Thorne collection is definitely from the Pacific drainage side. It is proba-
bly that this crest marks the southern limit of the unsilvered or partly
silvered form.
Reviewing, there is a mixture of unsilvered, partly silvered and
fully silvered callippe extending through the Southern Zone of Inter-
gradation, from the Greenhorn Mountains in the North, southward
through the Piute and Tehachapi ranges, to the crest of the series of
ranges extending transversely across California at about latitude 34°
40’ N. Most of the change from the predominantly unsilvered con-
dition in the north to the predominantly silvered condition in the
South, take place in about the northernmost one-fourth of the Zone.
Through the remaining three-fourths, beginning at about the mid-
length of the Piute Mountains, where a high degree of silvering al-
ready has been reached, the increase in silvering is very gradual. Al-
though it is still perceptibly incomplete at Lebec near the Southern
end of the Tehachapi, unsilvered and partly silvered become so rare
beyond there that one should not expect to encounter evidence of the
unsilvered condition without drawing large samples from the popu-
(i):}-20, ip62
VARIATION IN SILVERING
17
lation. It is reasonable to conclude that this is the southern limit of
the tendency toward lack of silvering in the ventral hind-wing spots
of callippe. To express this finding in Fig. 1, the area shaded with lines
slanted upward to the right should be extended to include localities
E, F, G and doubtfully H. When so extended, the southern limit of
lack of silvering would more nearly agree with the southern limits of
"uniformly colored red-brown upper surface” and "lack of brown pig-
ment between sdo<-s on the underside of the hind wings” as mapped
by Hovanitz (1943, Figs. 2 and 4).
DISCUSSION
Although this description adds some details to the distributional
pattern of silvering in callippe, it raises more questions than it an-
swers. Some are concerned with geographic limits. How much far-
ther north does silvering extend? Does it taper off gradually from
the western Greenhorn Mountains northward as does the opposite
condition from Walker Basin southward, or does it end abruptly a
short distance north of the sampled Greenhorn Mountain localities?
Does the more silvered population of the eastern slope of the Green-
horn Mountains end as in a cul-de-sac co-terminal with the limits of
the Kern River drainage, or does it extend northward as a tongue
along the higher Sierra Nevada toward silvered populations such as
those near Downieville and Gold Lake?
More perplexing questions concern the processes governing silver-
ing. The distribution of individuals among the seven score classes re-
veal a paucity of intermediates in the middle of the range (histo-
grams A and C in Fig. 2 ) . Could such a frequency distribution arise
from simple Mendelian inheritance, or is a more complex mechanism
required?
The existance of intermediate deo:rees of silvering suggests sim-
ilarity with the example of simple Mendelian inheritance described
by Ford (1945, p. 173) for the Currant Moth, Abraxas grossulariata
L., which, in its commonest form, has wings with a white bick.Erround.
In its less comomn form, lutea Cockerell, the white is replaced by a
deep yellow. When these two forms are interbred the heterozygotes
have a pale yellow background. Mating an individual homozygous for
white with one homozygous for deep vellov/, produces offspring inter-
mediate between the two parents. There is no dominance. Taking
this as a model for silvering in callippe, and using Ford’s system for
notation we may use the symbols and for represen-
ting the unsilvered ( Score 0 ) , the partlv silvered ( Scores 1 - 5 ) , and
the completely silvered (Score 6) phenotypes, respectively. For such
a model, in a population containing equal numbers of the two homo-
zygotes (C^C^ and OC^), we should find : C^C^:
1:2:1. None of our samples have equal numbers of homozygotes,
but if we pool samples A, B and the ratio is 24 : 51 : 18. This
18
SETTE
/. Res. Lepid.
differs from the ratio 1 : 2 : 1 by no more than one would expect
from random sampling variability in about half of the samples drawn
(P =z 0.44). We may conclude that there is no evidence against the
hypothesis that silvering is controlled by a single pair of alleles of the
phenotypes scored 1 to 5 may be lumped as the heterozygous com-
ponent of the population.
However, in the Currant Moth, equal doses for white and for
deep yellow produced a color about half way between the two homo-
zygotes. In callippe there are very few individuals about half silvered.
Individuals scored 3 and 4 comprise less than one-third of the cate-
gory with scores 1 to 5. Perhaps some more complex genetic system
may account for this peculiar distribution within the partly silvered
segment of the population. But I am attracted by the idea that some
chemico - physical process, such as crystallization may be involved.
These tend to be triggered by very slight differences in conditions,
and once triggered, go to completion.
In this connection it is also interesting that in areas of potential
silvering, the single scales, as viewed with a hand lense, appear either
totally buff or totally silver, never in between.
In pursuing this idea, I have found only a little information on
the silvery decorations of the Lepidoptera in the literature that bear
on its chemical nature. Mayer (1897) makes direct reference to its
nature in the Argynnids, saying ”... Dimmock ('83) has shown that
the silvery white and milk-white colorations are due to optical effects
produced by reflected light. In the silvery white scales, however, such
as the under surface of the hind wings of Argynnis, there must be
a reflecting surface toward the observor, for both silvery and milk-
white colors appear simple milk-white by reflected light.” According
to Fox (1953, p. 289) uric acid, derived from chrysalid metabolism,
deposited in the wings of the adult butterfly; guanine and uric acid
contribute opaque whiteness and glistening silvery aspects. Ford
(1945), although discussing extensively the physical and chemical
nature of butterfly coloration, is silent on the nature 'of silvering. Taylor,
(1925) points out that guanine (also spelled guanin) deposited in
crystalline form imparts the silvery appearance prevalently displayed
by pelagic fishes, while bottom fishes with prevalently white under-
surfaces have their "subdermal tissues heavily charged with amorphous
guanin, which is chalky white.” He describes guanine crystals, after
they have been processed into pearl essence for making artificial pearls,
as ". . . very thin light blades, floating in a liquid . . . [which] show
their maximum luster when they are oriented parallel to each other
. . . [when] pointing promiscuously in all directions, the effect will be
a metallic or dull pearly luster.”
While one would prefer evidence derived directly by analysis of
the substance as it occurs in callippe, it is not far fetched to suppose
that this substance is guanine, sometimes deposited as crystals disposed
t(i):3-20, i$6a
VARIATION IN SILVERING
19
with appropriate orientation to give the specular effect that we call
silvery, and sometimes in amorphous form adding whiteness to the
pigmental brown to produce the light buff of the unsilvered spots.
On this supposition it is necessary that the "silvering gene” control
only the conditions within the pupal scale-sac fluid so as to precipitate
guanine amorphously for buff scales, or alternatively, as crystals for
the silver scales. Presumably, in homozygotes the balance of influences
is well to one side of the critical point so that the deposite in all
scales is always amorphous, or always crystalline. In the heterozygotes,
too, it may take place on the "all or none” basis, but by single scales.
Under the equal and opposite influence of the two alleles the equilib-
rium may be very unstable and readily tipped to one side or the other
of the critical point. This could be mediated by very slight differences
in environmental conditions of the particular microclimate of a pupal
individual at the precise time of color deposition in the scale sacs.
This would tend to affect nearly all of the scales of an individual alike,
swinging most of them toward one side of the critical point more often
than nearly equal numbers to each side. Small differences between
members of the colony as to pupation site, date and time of day of
deposition of salts in the wing sacs, or even the rate of drying after
eclosion, might provide the variety of conditions necessary to produce
the observed peculiar frequency distribution of the supposed hetero-
zygous phenotypes.
This set of ideas is not advanced as an explanation, but as an
hypothesis inviting test by those having interest and competence in
the fields of experimental breeding or biochemistry, or both.
It is a pleasure to acknowledge the stimulus to undertake this
analysis and the material help received from William Hovanitz; the
generosity of Fred Thorne in giving specimens and sharing his uncom-
mon collecting serendipity; and the kindness of Paul Grey in review-
ing my identification of many hundreds of specimens of this and other
Argynnids.
LITERATURE CITED
ABRAMS, LEROY. 1940-1951. Illustrated Flora of the Pacific States, vols.
1-3, 4 vols. Stanford.
ABRAMS, LEROY and R. S. FERRIS. I960. Illustrated Flora of the Pacific
States, vol. 4, Stanford.
DOS PASSOS, C. F. AND L. P. GREY. 1947. Systematic Catalogue of
Speyeria with designation of types and fixation of type localities. Amer.
Mus. Novitates, no. 1370: 1-30.
JEPSON, W. L. 1923-25. A Manual of the Flowering Plants of California.
Berkeley. .
FISHER, R. a. 1948. Statistical Methods for Research Workers. Hafner,
N. Y. 10th Ed.
GUNDER, J. D. 1930. Butterflies of Los Angeles County, California.
Bull. Sou. Calif. Acad. Sci. 29 (2) ; 39-95.
HOVANITZ, WILLIAM. 1941. Parallel ecogenotypical color variation in
butterflies. Ecology 22: 259-284.
20
SETTE
J. Res, Lepid.
1943. Geographical variation and racial structure of Argynnis callippe
in California. Amer. Nat. 77:400-425.
MAYER, ALFRED G. 1897. On the color and color-patterns of moths and
butterflies. Bull. Mus. Comp. Uool. Cambridge 30 (4) : 169-256.
RICKER, W. E. 1940. Relation of "catch per unit effort” to abundance
and rate of exploitation. /. Fishery Res. Board Canada, Toronto.
TAYLOR, HARDEN F. 1925. Pearl essence: its history, chemistry, and
technology. Appendix to: Rept. U.S. Commissioner Fisheries for 1923.
Bur. Fish. Doc. 989. Washington, D.C.
1(1) :21.42, 1962
Journal of Research on the Lepidoptera
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Copyright 1962
THE EFFECT OF VARIOUS FOOD PLANTS ON
SURVIVAL AND GROWTH RATE OF PIERIS^
WILLIAM HOVANITZ AND VINCENT C. S. CHANG
California Arboretum Foundation, Inc., Arcadia^, Los Angeles State College,
Los Angeles and University of California, Riverside
The close relationship between most phytophagous insects
and their host plants is a general phenomenon known by most
entomologists. This relationship is so outstanding that field naturalists
can often locate colonies of any particular insect most readily by first
searching out the host plants, as they often are larger and more obvious
to the eye than the insects themselves.
What is not so well known is ( 1 ) the extent to which it is possible
for any particular species or race to survive on various food plants,
and (2) the causes of the attraction to, and survival of, any race of
insect on a particular plant.
It is the purpose of this paper to indicate something of the range
of ability to survive of two insects, Pieris rapae and Pieris protodice,
(the "cabbage or mustard "butterflies) on various cruciferous plants.
In other papers, it is intended to go into the various aspects of the
development of food plant preferences in insects, including its causes,
evolution, and relationship to origin of phylogenetic groupings.
THE MATERIAL AND METHOD
The experimental work to be reported on here and in subsequent
papers has been carried out in Arcadia, Southern California. All species
of the genus Pieris (Lepidoptera: Pieridae) found in southern Cali-
fornia have been utili2ed for the study. The geographical ranges and
specific food plants of the species existing in southern California are
being described elsewhere (Hovanitz, 1962) and the general distribu-
tion of the species of North America in another paper (Hovanitz,
1962). Five species of Pieris exist in this area, P. rapae (a European
immigrant), P. protodice, P. beckeri, P. sisymbrii and P. napi. Each
of these occupies a specific habitat and is restricted to a different food
plant.^
1 Aided by a grant from the National Science Foundation, Washington, D.C.
2The authors wish to express their great appreciation to Dr. W. S. Stewart, Director, Los Angeles
State and County Arboretum for great cooperation in making this work possible.
3Some taxonomists consider Pieris occidentalis to be a distinct species separate from Pieris protodice,
rather than a geographical race, ecological race or local phenotypic form. If this were the
case, there would be six species in southern California.
21
22
HOVANITZ AND CHANG
/. Re$. Lepid.
Pieris rapae is the easiest species to breed under greenhouse condi-
tions and has been used for most experiments. It aslo has a wider
range of satisfactory food plants than the others. As a European
immigrant, it remains restricted largely to plants of European origin
and has never adapted itself under natural conditions to any American
native. Contrariwise, Pieris protodice, a native species, has never
adapted itself to any European plant grown in cultivation, but has
assumed an "economic” role only in utilization of European plants
(mustards) growing in semi-wild habitats. The other Pieris are
restricted entirely to one or a few native species of plants.
The food plant tests were made in several series at different times
of the year and with different species of plants. Each series was run
for the purpose of comparing the influence of several plants (usually
five) on growth rate and mortality. For each series, a group of twenty
larvae, obtained as follows, were grown on each plant. Female Pieris
rapae were allowed to oviposit on a kale plant in the greenhouse. For
the first series, the eggs were removed from the kale leaf and placed
on leaves of the plant being tested in a petri dish. On all other tests,
the eggs .were allowed to hatch on kale, and to eat on it for one or
two days before removal to the tested plants. In order to minimize the
effects of variable environmental conditions, all tests of a series were
grown at the same time and place. For example, eggs laid on September
11, 1959, were used for the first test. On September 14, 1959, the
larvae were measured for the first time, and then successively thereafter
each day until pupation. The dates of larval (or pupal) deaths were
noted.
The later tests were similar except that (1) the origin of the
parents was different, ( 2 ) one or two day old larvae were used instead
of eggs, ( 3 ) the time of year and ( 4 ) place of breeding was different.
Data on origin of the material for each test are indicated in Table 1.
TABLE 1 : Source of larvae used in these experiments.
Series
Species
Food Source
Locaiity
( coilected
Date Generations
of experient from wild
Western
1
Pieris rapae
Cabbage
Orange
Sept.11,1959
1
County
2
Oct. 9,1959
2
3
Nov. 25, 1959
4
4
Feb. 20, 1960
6
5
Feb. 20, 1960
6
6
Aug. 2, I960
11
7
Sept. 19, I960
12
8
Pieris protodice
Mustard
Laguna Beach
Aug. 27, 1959
1
9
II
San Fernando
Valley
July 18, 1960
1
10
Cleome
Owens Valley
July 21,1960
1
i(t):2i-4S, 1962
EFFECT OF FOOD PLANTS
23
SERIES NO. 1
Series No. 1 was conducted for the purpose of comparing the
effect of the following plants on growth rate, size and mortality: black
mustard (Brassica nigra), garden nasturtium (Tropaeoum majus) ,
bladder pod (Isomeris arborea) and watercress {Nasturtium officinale)
( Table 2 ) . The females used for the source of larvae were obtained
from an open cabbage field isolated from any other cruciferous food
plant source and over which the adults were swarming in huge num-
bers. The probability that they came from any other source is
infinitesimal.
TABLE 2. Viability and growth rate of Pieris rapae grown on
different kinds of food plants ( Series 1 ) .
Plant
Number
Number and percent
of larvae died
Days from
egg to
pupation
Range of
variation in
larval growth
Nasturtium
20
16
30%
20
3
Kale
20
2
10%
12
2
Mustard
20
8
40%
13
3
Watercress
20
18
90%
17
1
Nasturtium
20
16
80%
18
5
Isomeris
20
10
50%
20
3
The mortality (Table 2. Series 1) was highest on watercress
(90%), next highest on garden nasturtium (80%), then Isomeris
(50%), mustard (40%) and least on kale (10%). The rate of
development as determined by the first larva to pupate, was most rapid
on kale (12 days), least rapid on Isomeris (20 days), and increasingly
more rapid on nasturtium (18 days) , watercress (17 days) and mustard
(13 days). The range in variation of larval growth was greatest for
nasmrtium, since five days elapsed between the first larva to pupate
and the last. The range as indicated in the table should be interpreted
with the fact in mind that only a fraction of the original twenty larvae
remained at the time of pupation. In other words, only two larvae
pupated out of twenty that were fed on watercress. Had more survived,
the range between the dates of pupation might have been greater.
The growth rates of the larvae on the various plants were obtained
by measuring ten larvae of each group each day and obtaining the
average length. These were then plotted and a curve drawn for each.
These curves are shown in Figure 1.
Measurements of the larvae were started on the third day after
hatching as they were so small prior to that time. However, great
■^It should be noted that the watercress used in these experiments was obtained by purchase in a
food market while all other plants were grown in the laboratory. It has occassionally been
found that despite repeated washings, insecticide residues may be present upon such commercial
vegetables and that these may cause death of larvae feeding upon them.
24
HOVANITZ AND CHANG
I. Res. Lepid.
differences already were apparent between the larvae grown on the
various plants. Those on kale were 4.7mm long as compared with those
on Isomeris which were only 1.5mm long. Those on nasturtium were
1.8mm, those on watercress were 1.9mm and those on mustard were
3.9mm. The kale-group larvae were over three times the size of the
Isomeris group larvae. Increase in size of the larvae on subsequent
days was fastest v/hen the larvae were grown on kale, second fastest on
mustard, and about equally as poor on each of the other three plants.
The largest sized larva was that grown on Isomeris (24mm) as com-
pared with that on kale (22.2mm) but after eight more days of larval
life. Those on watercress and nasturtium pupated at 20mm and those
on mustard at 17.9mm.
It appears that the greatest proportional size difference in the
larval groups occurred during the first three days, but that most of the
loss in size was later recovered by a longer growth period.
SERIES NO. 2
The tests of Series No. 2 were carried out in order to correct what
were thought to be defects in the experiments of Series No. 1. These
were two in number. The first was the high mortality in the eggs when
they were removed from kale and placed on the other plants. This
problem was averted by letting the larvae hatch on the kale plant before
moving to the tested plants. The second presumed defect was the
possibility of increased mortality of the very young larvae on the new
plants, due solely to physical characteristics, toughness, etc. This was
partially averted by allowing the larvae two days of feeding on kale
before transference to the tested plants.
One change in food plant was made in this test: radish obtained
from a food market was employed in place of watercress. Another
inevitable change was the climatic condition under which the tests
were conducted. They were changed slightly by the fact that the
tests were carried out starting October 9, 1959, instead of
September 11, 1959.
TABLE 3. Viability and growth rate of Pieris rapae grown
on different kinds of food plants (Series 2).
Days Range of
from variation
egg to in
No. and % No. and % Total No. pupa- larval
Plant No. of larvae died of pupae died and % died tion growth
Kale
20
2
10%
0
0
2
10%
11
2
Mustard
20
2
10%
3
15%
5
25%
13
4
Radish
20
8
40%
5
25%
13
65%
13
5
Nasturtium
20
8
40%
2
10%
10
50%
14
5
Isomeris
20
12
60%
3
15%
15
75%
15
3
The mortality (Table 3) in this series was generallly not so high
PIER IS RAPAE
Growth rates of larvae of Pieris rapae grown on various plants ( Series No. 1 ) .
26
HOVANITZ AND CHANG
/. Res. Lepid.
as in the series just preceding, though that on kale was the same at
10%, those on mustard dropped from 40% to 10%, on nasturtium
dropped from 80% to 40% and on Isomeris increased from 50% to
60%. Watercress was not tested this time but instead radish^ was used,
with a resulting mortality of 40%. The better survival of these larvae
is probably correlated with the fact that they were all grown during the
first two days on kale.
Generally, the time (minimum days) to pupation in this series
has been decreased (Fig. 2), The time on kale has been decreased
one day, on mustard it remains unchanged, and on nastrutium and
Isomeris it has been reduced five days. These reductions are probably
correlated with the more successful survival of the young larvae on
kale for the first two days, as compared with Series No. 1.
SERIES NO. 3
The tests of Series No. 3 are identical with those of Series No. 2
with the exception that they were started November 25, 1959 instead
of October 9, 1959; in addition, the larv.ae by this time were four
generations removed from the wild and inbred.
The larval mortality for all plants other than nasturtium was in-
creased considerably in Series No. 3 compared with Series No. 2;
10% to 30% for kale, 10% to 35% for mustard, 40% to 80% for
radish and 60% to 75% for Isomeris (Table 4). The data are com-
parable for the percentage of pupae died.
TABLE 4. Viability and growth rate of Pieris rapae grown
on different kinds of food plants ( Series 3 ) .
Days Range of
from variation
Plant
No.
No. and %
of larvae died
No. and %
of pupae died
Total No.
and % died
egg to
pupa-
tion
in
larval
growth
Kale
20
6
30%
0
0
6 30%
15
2
Mustard
20
7
35%
0
0
7 35%
16
5
Radish
20
16
80%
1
5%
17 85%
19
4
Nasturtium
20
7
35%
1
5%
8 40%
16
6
Isomeris
20
15
75%
2
10%
17 85%
18
8
The length of larval life has also been increased considerably in
Series No. 3 tests, and thus also the range of variation in larval growth.
The reasons for these differences between Series No. 2 and No. 3
are probably two fold. First, the later time in the season has meant
that the conditions of growth were somewhat different, primarily in
the temperature.
The larvae were grown in a greenhouse in which the temperature
was not well-regulated. Therefore, the temperature of the October
^The radish used in this particular experiment, but not in others reported on by us, was obtained
in a food market. Refer to footnote (4) for possible latent effects.
(mm) 3VAMV1 JO H10N31
(i):2i-42, 1962
EFFECT OF FOOD PLANTS
27
tests (Series No. 2) was higher than for the November tests (Series
No. 3). This would profoundly influence the results insofar as the
absolute growth rates are concerned. On the other hand, it did not
affect the relative growth rates on the various food plants. These are
still in the same descending order: kale, mustard, radish, nasturtium
and Isomeris.
0 ■ I 1 ■ ! I I ■ ■ U— I I
U 2x 3 4 5m 6 r 8 9x )0 ItxK !2 13 14 15
DAYS AFTER HATCHING FROM EGG UNTIL PUPATION NO OBSERVATION
MX = NO OBSERVATION ON KALE
2. Growth rates of larvae of Fieris rapae grown on various plants (Series
No. 2).
The differential increase in size of the larvae on the various food
plants is better shown in Series No. 3 (Fig. 3) than in Series No. 2
( Fig. 2 ) . This is because the slower development rate served to
exaggerate the differences between each strain. The larvae grown on
kale grew larger at a faster rate than the larvae on any other plant,
followed by mustard, nasturtium, Isomeris and radish. In Series No. 2,
the order of the latter two were reversed. It is interesting to note that
the ultimate size of the larvae which grew slowest was seldom equal
to the size of those larvae which grew fastest. In fact, there is almost
a direct relationship between these events.
A second reason for the difference between the mortality and
growth data of these two series may be of genetic significance. The
extent of inbreeding in the laboratory population by the fourth genera-
tion would have been sufiicient to (a) increase the mortality, (b) slow
20 0 1_ P/ERIS RAPAE
HOVANITZ AND CHANG
J. Res. Lepid.
28
O
(u/m) 3VAi/Vl JO H19N31
DAYS AFTER HATCHING FROM EGG UNTIL PUPATION (x=NO OBSERVATION)
(i):2i-42, 19^2
EFFECT OF FOOD PLANTS
29
development rate, (c) decrease size, (d) magnify the difficulties of
feeding on abnormal plants by the simple means of decreasing overall
vigor by the accumulation of allelic recessive lethal genes in the
laboratory strain which had not yet had a chance to be eliminated by
the selective effects of homozygosity.
SERIES NO. 4 AND 5
Series No. 4 was a test to compare various garden vegetables for
their usefulness to Pieris rapae. These vegetables were mustard, kale,
kohlrabi, broccoli and brussels sprouts. Series No. 5 was a similar test
for various other plants. These were mustard (as before), nasturtium,
radish, turnip and Isomeris. The tests were made in two series on
February 20, 1962, using larvae following six generations of inbreeding,
but of the same strain as used heretofore. In both of these tests, the
larval mortality was higher on comparable plants used in previous tests.
This again was probably due to the effects of inbreeding.
TABLE 5. Viability and growth rate of Pieris rapae grown
on different kinds of food plants (Series 4).
Days Range of
from variation
Plant
No.
No. and %
of larvae died
No. and %
of pupae died
Total No.
and % died
egg to
pupa-
tion
m
larval
growth
Mustard
20
9
45%
0
0
9
45%
15
9
Kale
20
5
25%
2
10%
7
35%
15
10
Kohl Rabi
20
9
45%
0
0
9
45%
14
9
Broccoli
20
9
45%
0
0
9
45%
14
9
Brussels
Sprouts
20
10
50%
0
0
10
50%
14
9
In Series No. 4 (Table 5), the data show that the larvae on kale
had the lowest mortality rate (25%) while all the other plant tests
were about the same (45-50%). Despite this, however, larvae from
kohlrabi, broccoli and brussels sprouts reached pupation ahead of those
on kale or mustard.
TABLE 6. Viability and growth rate of Pieris rapae grown
on different kinds of food plants ( Series 5 ) .
Plant
No.
No. and %
of larvae died
No. and %
of pupae died
Total No.
and % died
Days
from
egg to
pupa-
tion
Range of
variation
in
larval
growth
Mustard
20
9 45%
0
0
9
45%
15
9
Nasturtium
20
19 95%
0
0
19
95%
16
Radish
20
14 70%
2
10%
16
80%
15
13
Turnip
20
13 65%
0
0
13
65%
15
4
Isomeris
20
12 60%
0
0
12
60%
18
14
30
HOVANITZ AND CHANG
/. Res. tepid.
In Series No. 5 (Table 6), the data show that the lowest mortality
again was on mustard but with considerably higher mortality on the
other plants: for example, 95% on nasturtium, 70% on radish and
60-65% on turnip and Isomeris. The length of larval life was longest
on Isomeris (18 days) and about the same (15-16 days) on other
plants. The range of larval growth period varied greater with radish
and Isomeris than with any of the other.
4 5 6 7 ex 9 10 II 12 13 14 I5x . 16
DArS AFTER HATCHING FROM EGG UNTIL PUPATION tx^NO OBSERVATION)
4. Growth rates of larvae of Pieris rapae grown on various plants (Series
No. 4).
The rate of increase in size (Fig. 4) was more rapid with broccoli,
kohlrabi and brussels sprouts than with kale and mustard. It is believed
that this may be correlated with a more turgid or water-content condi-
tion of these plants as compared with the drier or stiffer condition of
the kale and mustard. For some reason, in Series No. 5, also, the
increase in size of the larvae was not up to expectations on the mustard
since the larvae on radish, turnip and nasturtium all exceeded those on
mustard in final size ( Fig. 5 ) .
SERIES NO. 6
Series No. 6 was tested on August 2, I960 with the use of larvae
eleven generations from the wild. One new plant, {Cleome lutea) , was
tested in this series, since this plant was found to be primary food
plant for Pieris protodice in Owens Valley, California. Mortality data
PiERiS RAPAE
i(i) 121-42, 1^62
EFFECT OF FOOD PLANTS
31
o o 00
in o in cJ
q
o
CVJ
(u/m) BVAyVl JO H10N31
Growth rates of larvae of Fieris rapae grown on various plants ( Series No. 5 ) .
32
HOVANITZ AND CHANG
/. Res. Lepid.
for kale , mustard, nasturtium and Isomeris, (Table 7) compared well
with previous data obtained on these plants (Series 1, 2, 3, 4 and 5).
Cleo7ne was only slightly less effective in maintenance of larval life,
the mortality rate being 30% as compared with 20% for kale and
mustard. The relative lengths of larval life and the range of variation
in larval growth also compares favorably with previous data. The
higher mortality as seen in Series 3, 4, 5 and 6 has here been reduced
to nearly the level of Series No. 1 .
TABLE 7. Viability and growth rate of Pieris rapae grown
on different kinds of food plants (Series 6).
Plant
No.
No. and %
of larvae died
Days from egg
to pupation
Range of
variation in
larval growth
Kale
20
4
20%
12
3
Mustard
20
4
20%
13
3
Nasturtium
20
10
50%
13
3
Cleome
20
6
30%
14
5
Isomeris
20
12
60%
18
6
The differences between growth curves for each of the plants
tested has been greatly increased in this test as compared with earlier
tests. The factors responsible for this are unknown but may be due
to a reduced adaptability caused by increased homozygosis (Fig. 6).
The larvae on kale were larger than those of any other plant at all
times during their growth period. Pupation occurred the twelfth day.
The larvae grown on nasturtium were almost the same size as those
grown on mustard, whereas in Series No. 1 (Fig. 1) the larvae on
mustard grew faster and larger. The larvae on Cleome were fourth in
size on any day, while those on Isomeris were fifth.
SERIES NO. 7
This series was conducted in order to place the plant Thelypodium
affine into position relative to the other standard test plants in the
laboratory. Thely podium affine is a native cruciferous plant from the
desert regions which is not known to be a food plant of any Pieris
species, though another Thely podium species is the food plant of Pieris
dsymhrii.
TABLE 8. Viability and growth rate of Pieris rapae grown
on different kinds of food plants ( Series 7 ) .
Plant
No.
No. and %
of larvae died
Days from egg
to pupation
Range of
variation in
larval growth
Kale
20
5
25%
16
3
Mustard
20
6
30%
17
4
T. affine
20
14
70%
22
5
Isomeris
20
12
60%
18
6
Nasturtium
29
8
40%
17
6
PIER IS RAPAE
■{1)121-42, 1962
EFFECT OF FOOD PLANTS
33
\
o
o
CSJ
fluuij 3VAHW1 JO HI9N31
DAYS AFTER HATCHING FROM EGG UNTIL PUPATION
34
HOVANITZ AND CHANG
/. Res, Lepii.
The tests of Series No. 7 (Table 8) were carried out on
September 19, I960, using larvae after twelve generations of laboratory
inbreeding and growing on kale. The larvae grew relatively well on
mustard, kale and nasturtium. The effectiveness of Isomeris was as
previously tested. Thely podium affine was the poorest of all these
plants as far as effectively providing the larva with stimulation to feed.
The mortality of the larvae on T, affine was 10% compared with
60% on Isomeris, 40% on nasturtium, 30% on mustard and 25%
on kale. The length of larva life was increased from 16 days on kale
to 22 days on Thelypodium. The final size of the larvae was reduced
from 22mm long on kale to only 18mm long on the Thelypodium
larvae which survived. (Fig. 7).
GENERAL OBSERVATIONS ON PIERIS RAPAE TESTS
The seven series of tests on Pieris rapae larvae have indicated
several significant points:
( 1 ) Larvae from strains grown previously on cabbage or kale
have a lower mortality rate, have a faster development rate, and a
greater size when grown on kale than when grown on any other plant
tested. When similar larvae are grown on mustard, these factors are
decreased slightly. The order of decrease for plants other than these
two is in the order approximately as follows: nasturtium, Cleome,
radish, Isomeris, and T, affine. Plants of the cabbage group: kohlrabi,
broccoli and brussels sprouts, are about the same as cabbage or kale.
Turnip is about the same as radish.
( 2 ) Higher mortality in the tests is generally correlated with a
slower development rate, a delayed time to pupation and smaller size
of the larva and pupa. There are some exceptions to this rule; for
example, the final size of the larva may be as great when the growth
rate was slow on one plant as when it was fast on another. This has
been indicated, however, only for plants of the cabbage group.
(3) Mortality rates in the various test series have been variable.
These variations in mortality have been thought not only to be due to
the genetic effects of inbreeding but also partly to the changed environ-
mental conditions encountered while the tests were conducted, such as
lower temperatures, etc. of the changing seasons. Lower temperatures
slow the developing rate, and increase the relative humidity. Increased
humidity permits the larvae to be attacked by bacterial or viral diseases
to a much greater degree.
SERIES NO. 8
Tests of Series No. 8, 9 and 10 were conducted with larvae of
Pieris proto dice rather than with Pieris rapae.
The first series with this species (No. 8) was conducted on
August 27, 1959, utilizing larvae one generation from the wild. The
fwu/} 3VAHV1 JO H10N31
DAYS AFTER HATCHING FROM EGG UNTIL PUPATION ( NO OBSER V ATION
36
HOVANITZ AND CHANG
/. Res. Lepid.
females of the previous generation were obtained in a field of black
mustard, not near any other source of food plant, and in which eggs
and larvae were found on that plant. This is good' evidence that the
natural plant utilized here as larval food was mustard.
In this series, the food plants tested as larval food were mustard,
kale, radish, Isomeris and nasturtium. The mortality of larvae of
Pieris protodice on these plants increased in the order just stated,
namely, 10% on mustard, 20% on kale, 55% on radish, 55% on
Isomeris and 95% on nasturtium (Table 9). Deaths in the pupa stage
on kale, radish and Isomeris materially increased the total mortality
before adult emergence. This was not increased on mustard.
TABLE 9. Viability and growth rate of Pieris protodice grown on
different kinds of food plants (Series 8).
Plant
No.
No. and %
of larvae died
No. and %
of pupae died
Total No.
and % died'
Days
from
egg to
pupa-
tion
Range of
variation
in
larval
growth
Mustard
20
2
10%
0 0
2
10%
10
2
Kale
20
4
20%
2 10%
6
30%
11
2
Radish
20
11
55%
4 20%
15
75%
13
3
Isomeris
20
11
55%
3 15%
14
70%
16
3
Nasturtium
20
19
95%
■— — -
19
95%
21
The length of larval life was greatly different on the various food
plants, ranging from ten days on mustard, eleven days on kale, thirteen
days on radish, sixteen days on Isomeris to twenty-one days on nastur-
tium. This wide variation in length of larval life is greatly different
from anything encountered with Pieris rapae. It would indicate a
fundamental difference in food plant adaptability between these two
species, Pieris rapae being much more versatile in its requirements than
Pieris protodice.
The size of the larvae at various days after hatching (Fig. 8)
indicates that the mortality data are in close correlation with the size
increments of the larvae with respect to food plants involved. Larvae
on mustard and kale increased to 24mm in length, those on radish,
Isomeris and nasturtium to 20-2 1mm each.
SERIES NO. 9
Discovery that Pieris protodice utilizes Cleome lutea as a natural
food plant in the Owens Valley prompted a series of tests involving
this plant. Both Series No. 9 and Series No. 10 were carried on for
the purpose of testing Cleome in comparison with other plants used
as standards. Series No. 9 utilized Pieris protodice larvae bred from
adults obtained from mustard fields in the San Fernando Valley while
Series No. 10 utilized Pieris protodice larvae obtained from Cleome
pf£ms pporoDicE
i(i);2i-42, i$62
EFFECT OF FOOD PLANTS
37
mrs AFTER HATCHING FROM EGG UNTIL PUPATION (NO observation = x j
38
HOVANITZ AND CHANG
/. Res. Lepid.
plants in the Owens Valley (Big Pine). There is no chance that the
San Fernando population had access to Cleome but there is a chance
that the Owens Valley population did have limited access to mustard.
TABLE 10. Viability and growth rate of Pieris proto dice grown on
different kinds of food plants (Series 9).
Plant
No.
No. and %
of larvae died
Days from egg
to pupation
Range of
variation in
larval growth
Mustard
20
6
30%
14
3
Cleome
20
7
35%
14
4
Nasturtium
20
18
90%
17
2
Isomeris
20
8
40%
17
5
The Series No. 9 larvae (Table 10) showed highest mortality on
nasturtium (95%), just as in Series No. 8, with the lowest mortality
on mustard (30%), next highest on Cleome (35%) and then Isomeris
(40%). This corresponds well with comparable data in Series No. 8.
The growth increment curves for this series are somewhat com-
parable with the preceding series except for the one surviving larva
on nasturtium which increased to a size greater than that attained by
larvae on any other plant. The differences between the growth curves,
however, are not as distinct and clear as in Series No. 8, perhaps because
of temperature differences between the two. Cleome is almost as
satisfactory as mustard as a food plant for these larvae. ( Fig. 9 ) •
SERIES NO. 10
This series was run almost contemporaneously with the preceding
and involved larvae from the Owens Valley strain grown on Cleome.
The mortality (Table 11) was lowest on mustard (20%), second
TABLE 11. Viability and growth rate of Pieris protodice grown on
different kinds of food plants (Series 10).
Plant
No.
No. and %
of larvae died
Days from egg
to pupation
Range of
variation in
larval growth
Mustard
20
4
20%
11
3
Cleome
20
6
30%
12
5
Isomeris
20
7
35%
13
7
lowest on Cleome (30%) and then Isomeris (35%). There was a
greater range of larval growth rates on this group of larvae on Isomeris
than in any other group as well as the lowest mortality on this plant
in any of the larvae tested. The growth increment curves for these
larvae indicate that the larvae do not show much difference in their
reaction to mustard and Cleome but that mustard is still slightly better,
and Isomeris is always poorer. Final size of the larvae, however, was
larger for the Cleome (23 mm) than for the mustard (22 mm) or
the Isomeris (18 mm)'. (Fig. 10).
LEN6TH OF LARVAE (mm) LENGTH OF LARVAE (>
i(i):2\-42, 1962
EFFECT OF FOOD PLANTS
39
DAYS AFTER HATCHING FROM EGG UNTIL PUPATION
9. Growth rates of larvae of 'Pieru protodice grown on various plants (Series
No. 9).
PAYS AFTER HATCHING FROM EGG UNTIL PUPATION
10. Growth rates of larvae of Pieris protodice grown on various plants (Series
No. 10),
40
HOVANITZ AND CHANG
/. Res. Lepid.
GENERAL OBSERVATIONS ON PIERIS PROTODICE TESTS
The three series of tests on Pieris protodice have indicated several
significant points: (1) Larvae from strains obtained from mustard
or Cleome populations had equally good survival value on mustard
plants. The following plants gave decreasing survival value to the
larvae: (Series 8) kale, radish, Isomeris and nasturtium. (Series 9)
Cleome, nasturtiuna, Isomeris. (Series 10) Cleome, Isomeris. (2) Higher
mortality in the tests is generally correlated with a slower development
rate, a delayed time to pupation and smaller size of larvae and pupae.
DISCUSSION
The tests described in the preceding pages have indicated some
significant information regarding the effect of various food plants on
survival in two species of Pieris. The ability to survive on different
food plants is not an all or none relationship, but rather is a relation-
ship based upon relative ability to survive. In the most extreme case,
all larvae may die rather than eat a completely unacceptable plant.
On the other hand, plants may be placed in an order of desirability
with regard to the ability of larvae to survive, or to grow satisfactorily,
on plants which they may accept at least reluctantly.
The tests described in this report are based upon using three criteria
to indicate the desirability of a plant to the larvae. The first of these
is the ability to survive, or the reverse which is the mortality, expressed
as a percentage of the total larvae which died during the experiment.
The second criterion is the daily size increment during the life of the
larva. This actually tests two factors: (1) the physiological usefulness
of the food to the larvae and (2) the total amount of food eaten and
uilized, which may be a direct result of the stimulatory activity of the
plant toward the larvae. The third criterion is the range in variation
of larval growth. The significance of this criterion is in more doubt
than the other two, but the range indicated would in some degree show
the genetic variability present in the larval population toward physio-
logical adaptation to the particular plant.
The results just described show that some plants are much more
suitable for larval food to certain larvae than are other plants. The
reasons for this greater suitability are not made clear. In those cases
where higher mortality, slower growth rate, size increments and final
size are correlated, the cause of the poor growth could be lack of
nutriment in the plant, or lack of larval ingestion of the plant. In other
cases where larval mortality may be high, as with the broccoli, Kohl-
rabi and Brussels sprouts tests (Table 5) hut growth rate rapid and
size large ( Fig, 4 ) , there are probably two factors involved. The rapid
increase in size could be due to a high attraction to a very nutritious
plant, but secondary conditions surrounding the plant may be conducive
I (i):2i -42, 1962
EFFECT OF FOOD PLANTS
41
to a higher mortality, namely, high water content in the plant itself,
or higii humidity surrounding tne plant whicn would increase the
susceptability of the larvae to virus and bacterial intections.
Toughness of leaves of a plant cannot be ruled out as a factor
involved in larval speed of growth. Some leaves are too tough for the
larvae to feed upon easily. Leaves of the same plant vary in tnis regard;
succulent leaves at the siioot apex are usually much more desirable than
older, mature leaves. This may be caused by toughness, or it may be
caused by the greater perception of attractive substances permeating
from younger leaves. At any rate, the relative uniformity of the tests
in the difterent series is good evidence for the conclusion that this
factor has not been of major significance.
The greater ability of tieris rapae as compared with Pieris protodice
to survive on a variety of common cruciferous plants is interesting in
view of the fact that both succeed well on black mustard. Pieris rapae
survives better on kale over mustard and P. protodice survives better on
mustard over kale. However, on most other cruciferous plants tested,
there was a wide disparity between their preferences. The selections
made by the strains of larvae shown by these experiments complements
perfectly the known differences in geographical and ecological distribu-
tion of the two species concerned, and indicates that there is little
natural competition between the species for food plants. This shows
also why Pieris rapae is always associated with European cruciferous
plants, weeds or truck crops, while Pieris protodice is generally
associated with other cruciferous plants.
Takata (1957, 1959, 1961) has published a number of papers
which bear upon this problem. None of these deals precisely with
the effect of various food plants on survival and growth rate of Pieris,
but rather deals only with selection preferences by the larvae and the
adult. Further considerations of the relation of this work of Takata
on Pieris rapae crucivora to the present general program will therefore
be made in a complementary paper entitled "Three Factors affecting
larval choice of food plant” to be published in this journal.
A visit to our project by W. H. Dowdeswell of England during the
course of these experiments was followed by a similar study of more
limited extent on Pieris napi L. in England. Dowdeswell found that
two populations of this Pierid were feeding on different cruciferous
plants. By switching the food plants in laboratory experiments on test-
ing the larvae, he found as we had, that there was a higher mortaity
and decreased growth rate when the larvae were fed on plants not
the normal food plant for that population (Dowdeswell and Willcox,
1961).
SUMMARY
1. Tests were made of the ability to survive of larvae of two species
of Lepidoptera, Pieris rapae and Pieris protodice on various food plants.
42
HOVANITZ AND CHANG
/. Res. tepid.
2. Three criteria were tested: (a) mortality, (b) growth rate
and size increment, (c) range in variation of larval growth.
3. Seven series of tests involving Pieris rapae were conducted
showing the results summarized under the heading "General Observa-
tions on Pieris rapae Tests.”
4. Three series of tests involving Pieris protodice were conducted
showing the results summarized under the heading "General Observa-
tions on Pieris protodice Tests.”
5. Pieris rapae has significantly different requirements for food
plants than Pieris protodice, even though the tested larvae were both
from mustard-bred strains.
6. The native adaptability of Pieris rapae to survive on various
plants is better than that of Pieris protodice.
7. The causes of differences in survival value of the larvae on
different food plants are believed to be several, but the main one is
the total amount of food eaten and utilized.
LITERATURE CITED
DOWDESWELL, W. H. AND H. N. A. WILLCOX. 1961. Influence of the
food plant on growth rate and pre-imaginal mortality in the green-veined
white butterfly Pieris napi (L). Entomologist 94:2-8.
HOVANITZ, W. 1962. The ecological and geographical distribution of
Pieris species in Southern California. J. Res. Lepidoptera 1 : in press.
HOVANITZ, W. 1962. The distribution of the species of the genus Pieris
in North America. J. Res. Lepidoptera 1 ( 1 ) : this issue.
TAKATA, N. 1957. Studies on the host preference of cabbage butterflies
{Pieris rapae L.) III. Mechanism of food preference of cabbage butterfly
larvae. Jap. J. Ecol. 7 (3) :117-119.
TAKATA, N. 1959. Studies on the host preference of common cabbage
butterfly, Pieris rapae crucivora Boisduval. IV. Tendencies of food prefer-
ence in relation to total quantity of the host plants cultivation. Zoological
Magazine (Japan) 68(5) : 187-192.
TAKATA, N. 1959. Studies on the host preference of common cabbage
butterfly, Pieris rapae crucivora Boisduval. VI. Change in the food
preference of larvae when reared successively by the definite food plant
for several generation. (Preliminary report) Jap. J. Ecol. 9:224-227.
TAKATA, N. 1961. Studies on the host preference of common cabbage
butterfly, Pieris rapae crucivora Boisduval. XI. Continued studies on the
oviposition preference of aduir butterflies. Jap. J. Ecol. 11 (3) :124-133.
TAKATA, N. 1961. Studies on the host plant preference of the common
cabbage butterfly, Pieris rapae crucivora Boisduval. XII. Successive
rearing of the cabbage butterfly larva with certain host plants and its
effect on the oviposition preference of the adult. Jap. J. Ecol. 11(4):
147-154.
Journal of Research on the Lepidopvera
l(l):43-49, 1962
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Copyright 1962
GENERAL CHARACTERISTICS OF THE
MOVEMENTS OF VANESSA CARDUI (L.)
J. W. TILDEN
San Jose State College, San Jose, Calif.
It is a matter of common knowledge that the populations of
many species of butterflies participate in large and extensive mass
movements. While several workers, notably C. B. Williams of England,
have done extensive work in this field, relatively few investigators
have made it their principal subject of research. Though much
pertinent information has been assembled and certain tentative con-
clusions may be reached, it cannot be said that the movements of
butterflies are as well understood as certain other phases of the study
of these insects.
Most movements of butterflies, while usually referred to as migra-
tions, would not meet the criteria set up by students of vertebrates for
defining migrations. The migrations of birds, for example, involve
cyclic movements. There is a going out and a return. Few butterflies
other than the Monarch [Danaus plexippus (L.)], habitually possess
such a rhythmic pattern of movement. Even with the Monarch, the
individuals that return are usually (probably always) not the same
individuals that initiated the outward movement, but are the offspring
of these original individuals.
Many movements of butterflies are relatively local wanderings, and
may be in one direction in one local area, but in a different direction
in another area a relatively few miles away. For example, in the fall of
I960, Tilden noted Libytheana hachmanii (Kirt.) (the Snout Butter-
fly) moving southwesterly in large numbers in the vicinity of Con-
tinental, Pima Co., Ariz., on Sept. 14. The next day, Sept. 15, equally
large numbers of this species were observed moving up-canyon ( north-
ward) in Sycamore Canyon, Santa Cruz Co., about thirty -five miles
south of Continental by air line, though considerably further by road.
The very conspicuous movements of certain species have been
regarded as depending on population pressures, leading to a mass
exodus. These may be considered as emigrations from the populated
area, and as immigrations into surrounding area. Such movements
frequently appear to be in one direction only. Unless the insects were
to find suitable conditions at the far end of the movement, establish-
43
44
TILDEN
/. Res. Lepid.
merit would not result. Such movements have been thought to account
in part for the finding of strays far beyond the normal ranges of
the species.
Movements of an altitudinal nature have been observed. Nymphalis
calif ornica (Bdv.) has been seen ascending the Sierra Nevada of
California in great numbers. Such movements may result in finding
sufficient larval food plants for a subsequent brood. In I960, Tilden
found this species in outbreak numbers of adults at Manzanita Lake,
Lassen Volcanic National Park, Calif., on June 10-17. Numerous larvae
in various stages of development were found on Ceanothus spp. Adults
could be observed many miles below the Park, working their way up
the mountain. It may be significant to note that this species passes
the winter as an adult. It seems plausible that these altitudinal move-
ments of adults are the usual way for adults to reach the higher eleva-
tions, where they act as parents of a late high elevational brood. This
might be a fruitful field of investigation.
The often spectacular "migrations” of Vanessa cardui (L.) present
an interesting condition somewhat intermediate between true migra-
tions and other types of movements. Return movements seem not to
be clearly documented in regard to this species. However, reproduction
has been observed repeatedly along the route, and the offspring take
up the movement when they emerge as adults. These mass movements
of Vanessa cardui thus proceed in periodic waves, alternating with
relative lulls between, and the whole process has much more continuity
than is characteristic of the movements of most other butterfly species.
Much of the basic work on the movements of Vaness-a cardui has
been done by C. H. Abbott. Others, including Sudgen, Woodbury and
Gillette in Utah, have made contributions. Among the various titles
that form the literature on the movements of this species in western
United States, some are short observational notes only. Others present
a considerable amount of detailed study. Abbott has attempted to
understand the nature of these movements and has produced several
penetrating papers on the subject.
The last great movement was in 1958. Abbott traces three or
more generations extending up to or beyond the Central Valley of
California. Other generations carried the waves through the Bay
Region of California on into Oregon. The generations involved in
the northern portions of the movement become less clear.
The first generation seems to have originated in northern Mexico,
the second in the Imperial Valley of California, western Arizona and
the eastern Mohave Desert of California, the third in western Mohave
Desert, and the fourth in the Central and Coastal Valleys of California,
and to the Bay Region. Abbott (1959) has presented material that
includes the observations of a number of workers, and has coordinated
this material. In addition, the follov/ing observations are here presented
for the first time.
j(i):4}-49, 1962
MOVEMENTS OF VANESSA
45
Tilden found many dead adults in western Arizona (Mohave
County) and in the vicinity of Needles, California, as well as large
numbers of moving adults, March 30-31, 1958. From Barstow, San
Bernardino Co., west to Mohave, Kern Co., great numbers of adults
were flying northwesterly on March 31. Later that day adults were
seen moving through Tehachapi Pass, Kern Co., and spilling into the
San Joaquin Valley east of Bakersfield, Kern Co. On April 20 to
May 9, eggs and larvae were abundant in the Salinas Valley,
Monterey Co.
On May 30 - June 1, Tilden found large numbers of adults along
the Redwood Highway (Route 101) between San Rafael, Marin
County, and Cloverdale, Sonoma County, moving northwesterly. Dead
adults were seen as far west as Booneville, Mendocino County, where
but a few living adults remained. This may mark the western edge
of the movement.
In the vicinity of The Geysers, Sonoma County, adults were moving
northwesterly in large numbers in overcast weather (fog) on June 1,
before noon. Larvae were abundant on Amsinckia (Fiddleneck) and
on the introduced Spanish Thistle ( Carduus ) , in various stages of
development.
Two interesting reports were from Oregon. Ray Albright reported
large numbers of Vanessa cardui moving in a north-north-easterly
direction on May 18-19, 1958, while on May 20-21, equally large
numbers were observed moving in exactly the opposite direction. He
attributes this reversal of direction to the insects meeting a cold front,
since a storm was in progress some distance beyond the place toward
which the insects were noted to be flying. The other report was from
David Huntzinger at Crater Lake National Park. He noted that Vanessa
cardui was abundant there in 1958, though scarce or absent in 1957.
On April 1, 1958, Tilden observed what is apparently a new
observation for the United States. Large numbers of Vanessa cardui
were flying north-westerly in the rain. Between Vidal, San Bernardino
County, and Desert Center, Riverside County, the rain was light to
moderate, but rather cold. Between Indio and Whitewater Canyon,
later the same day, the flight still continued in heavy to very heavy
rainfall. This seems to indicate that adults may continue to fly, once on
the wing, even in heavy precipitation. It would seem a priori that
initiation of a flight during rain might be less likely.
From observations so far recorded it is possible to generalize on
some of the characteristics of the movements of Vanessa cardui. Some
of these generalizations have been set forth by Abbott and others,
while some are suggested here for the first time.
1. The adults tend to fly into the wind. This may be a key point
in attempting to explain the prevailing direction of the movements.
During the time of year when Vanessa cardui is moving, the prevailing
winds in California are north-westerly, and the movements of the
46
TILDEN
/, Res. Lepid.
butterflies are also in a north-westerly direction, almost or quite directly
into the prevailing winds. Sudgen, in 1937, noted a northerly direction
of flight in Utah. Sudgen, Woodbury and Gillette, in 1947, noted a
north-easterly direction of flight. Observations to be made in the future
might very well take special note of the wind direction as well as the
direction of the insects’ flight, to find how complete this correlation
between wind direction and flight direction may be.
2. The adults move at a rather equal velocity in relation to one
another, resulting in a fairly constant spacing of the individuals.
There have been some observations made where this did not seem to
be the case, but for the most part, this generality seems to hold.
3. The insects tend to fly over obstacles rather than around them.
At times this appears to the observer quite amusing, even ridiculous.
If a tree, building or parked car obstructs the route, the adults fly over
the top of the obstruction, when in some instances it would appear
that a veering to the right or left would be more easily accomplished.
The obstruction, whether it be low or tall, is barely cleared. On
March 30, 1958, Tilden observed them piling directly over a small
building near Barstow, San Bernardino County, the only building for
miles in an open desert.
4. The adults tend to fly at rather even densities. If a line is
marked off between tv^o points, fifty or one hundred feet apart, and the
insects crossing this line per minute are counted, the results from
several one minute counts are surprisingly similar. If then, the observer
drives his car several miles, sets up another station and makes another
series of counts, the results will again be similar, and frequently quite
close to those of the previous station. But in widely separated areas,
the count may be very different. That is, while the density on the
Mohave Desert, for instance, may be quite constant, counts made in
San Diego County at another time might indicate a different density.
Further study may shed light on the possibility that density of flight
is related to density of population.
5. Vanessa cardui adults will fly in overcast, at least after the
movement is under way. This has been observed between Victorville
and Needles, San Bernardino County, March 31, 1958, and near The
Geysers, Sonoma County, May 31 and June 1, 1958.
6. They have been observed once at least in the United States
(vide supra) to fly in large numbers during moderate to very heavy
rainfall.
7. The migrating individuals are all of one species. Other species
of butterflies do not seem to become involved in the mass movements
of Vanessa cardtii.
8. The individuals in the flights are of both sexes. This was noted
independently by sampling in 1958, by both Tilden and O. E. Sette.
The sex ratio of the insects when engaged in a flight is very nearly
the same as that oi Vanessa cardui at any other time, 0.5 or one half
i(i):43-49> 1962
MOVEMENTS OF VANESSA
47
of each sex. Tilden took a sample of fifty adults, and found twenty-
seven males, twenty-three females. The numbers taken by Sette are
not recorded. This phase of the investigation deserves further study
and this is planned for the future.
9. The females from the above sample were not greatly gravid.
None had eggs ready to oviposit. This suggests that the females may
drop out of the flight when ready to oviposit. If this be true, it might
explain the tendency to reproduce along the flight routes.
10. The total movement proceeds in waves, or phases, which so
far appear to be based on broods, a conclusion which seems to be well
supported by the work of Abbott.
In the light of current findings, it is possible to reconstruct, at
least to some degree, the nature of the populations of Vanessa cardui
during and prior to the flights.
1. The first (initiating) brood apparently originates to the south,
in northern Mexico or southwestern Arizona.
2. Findings from those movements that have been studied indicate
that large flights can occur only in years when there has been sufficient
rainfall on the deserts for a large growth of vegetation, enough to
support such populations of Vanessa cardui.
3. The food plants in the desert areas, as noted by many observers,
are principally Boraginaceae, especially Cryptantha, but also Amsinckia.
Malvaceae have also^ been reported. Thistles and other composites seem
to be of little importance at this time.
4. Upon emergence of the adults, each succeeding brood begins
to fly into the wind, which in the deserts of California at this season
(early spring) results in a north-westerly direction of flight.
5. These adults reproduce at some point along the line of flight.
Time and place of reproduction presumably depend on the condition
of the insects and the presence of suitable vegetation. Information on
how far the females fly before reproducing, and the precise reason
for selecting a certain place for reproduction, would add much to our
knowledge.
6. Four to five broods, perhaps more, succeed one another before
the force of the movement is spent.
7. The northern extent of the movement is at least to the San
Francisco Bay Region of California. In some years, such as 1958, the
effect extends into northern California, and apparently further, to
Oregon and perhaps even Washington. Information as to the relation-
ship between the Vanessa cardui in California and those in Oregon
and Washington during such outbreak years is badly needed.
Williams and Abbott have raised the question as to whether
Vanessa cardui is found north of the Imperial Valley, Imperial County,
Calif., in any but outbreak years. There seems to be good evidence
that it is. An examination of pinned specimens from many parts of
California shows specimens taken in almost every year for many years
48
TILDEN
/. Res. Lepid.
back. As a Professor at San Jose State College, Tilden has examined
hundreds of student collections over a period of fourteen years, and
has also examined other specimens dating back to 1920. Some speci-
mens of Vanessa cardui appear in student collections every semester.
It is evident that this species is a normal component of the butterfly
fauna in many parts of California. It is probable, however, that this
local endemic population has little or nothing to do with the mass
movements which from time to time are superimposed on it.
Two other items seem worth mentioning. Firstly, moderate to
large populations of Vanessa cardui may be found late in the fall at
high elevations in the mountains of the western United States. Specifi-
cally, Tilden found hundreds of adults at Hannagan’s Meadows, White
Mountains, Arizona, September 12, I960. A fair population was
found at Barton Flats, San Bernardino County, Calif., in September,
1957. Similar populations have been reported from the Sierra Nevada
of California and elsewhere. The nature of these populations is not
known, but it would seem that they are independent of the populations
concerned with the great flights that occur in certain years.
Secondly, the mass flights of Vanessa cardui are of a very irregular
nature. They are not regularly cyclic and do not occur at predictable
intervals. However, some success in predicting these flights has been
possible through knowledge of rainfall on the desert. Several workers
foresaw the 1958 outbreak in the gathering populations of the south-
west deserts.
An interesting and provocative suggestion of how rainfall may be
necessary for mass movements of this species, was observed by Tilden
September 11, I960, at Yucca, Mohave County, Arizona. Late summer
rains were plentiful and vegetation was well developed. Numerous
freshly emerged adults of Vanessa cardui were present, and reproduc-
tion was in progress, with larvae of various stages on Cryptantha. It
seemed that the basis for a movement was all prepared. However, the
winter rains did not materialize, and on March 26, 1961, the area
around Yucca was dry and no Vanessa cardui were to be found. If
this abortive season is contrasted with the abundant rainfall of 1958
over the entire area, it points to a dependence of these populations
on ample winter rainfall in the areas that are far enough south to act
as reservoirs for populations that can engage in a mass flight.
Several questions are raised : ( 1 ) Why do these insects move at
all, rather than remaining where they are? Possibly food may become
^scarce, but populations exist that do not move out in such a systematic
manner under lowered food conditions. (2) Why is the flight directed?
Why do not these insects move randomly in any direction open to
them? The tendency to fly into the wind may be a clue here. Or have
such movements been repeated so many times over the history of the
species, that the tendency to fly in one direction has been selected for?
(3) What causes the adults to reproduce here and there along the
(i):43-49< ^9^2
MOVEMENTS OF VANESSA
49
route? Why do they not merely fly on until exhausted? Present
information may suggest some tentative answers, but such answers,
while plausible, are not yet supported by suflicient data.
The author would add that this attempt at a synthesis of our
knowledge concerning the movements of Vanessa cardui in western
United States is prompted more by interest on his own part than by
a desire to inform others. The purpose of this paper is to bring into
focus our ignorance of the real basis of this interesting phenomenon,
with a hope that future work will help to clarify some of the little
known facets of the problem.
LITERATURE CITED
ABBOTT, C. H. 1941. The 1941 migration of the painted lady butterfly,
{Vanessa cardui) in southern California. Bull. Ecol. Soc. Amer., 22:13.
— — — 1946. Mapping the 1945 migration of Vanessa cardui in southern
California. Bull. Ecol. Soc. Amer. 27 :49.
— — 1950. Twenty-five years of migration of the painted lady butterfly,
Vanessa cardui, in southern California. Pan-Pac. Ent., 26:616-172.
— - 1951. A quantitative study of the migrations of the painted lady
butterfly, Vanessa cardui L. Ecol. 32:155-171.
— — 1959. The 1958 migration of the Painted Lady Butterfly, Vanessa
cardui (Linnaeus), in California. Pan-Pac. Ent., 35:83-84.
ESSIG, O. E. 1926. A butterfly migration. Pan-Pac. Ent., 2:211-212.
SMITH, RAY F., & E. GORTON LINSLEY. 1945. Migration of Vanessa
cardui (Linn.). Pan-Pac. Ent., 21:109.
SUGDEN, JOHN W. 1937. Notes on the migrational flights of Vanessa
cardui in Utah. Pan-Pac. Ent., 13:109-110.
SUGDEN, JOHN W., ANGUS M. WOODBURY & CLYDE GILLETTE.
1947. Notes on the migration of the painted lady butterfly in 1945.
Pan-Pac. Ent., 23:79-83.
WILLIAMS, C. B. 1958. Insect Migration, xiii -f235 pp., 11 plates, 22
photos, 49 maps & diagrams. Collins, London.
WOODBURY, ANGUS M., JOHN W. SUGDEN & CLYDE GILLETTE,
1941. Notes on migrations of the painted lady butterfly in 1941. Pan-Pac.
Ent., 18:165-176.
Journal of Research on the Lepidoptera
1(1) :51-61, 1962
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Copyright 1962
THREE FACTORS AFFECTING
LARVAL CHOICE OF FOOD PLANT’
WILLIAM HOVANITZ AND VINCENT C. S. CHANG"
California Arboretum Foundation, Inc., Arcadia,
Los Angeles State College, Los Angeles and
University of California, Riverside
Prior to the initiation of comprehensive tests of the attraction
by larvae of Pieris rapae of various food plants, a number of prelim-
inary trials were made to test the effect of external factors which might
influence the correctness of these tests. Of the factors which were
thought most important, the following were selected for trial:
( 1 ) Influence of direction of air movement ( "wind”) .
(2) Influence of age (size) of larvae.
(3) Influence of food previously eaten by larvae, or strain of
larvae.
EFFECT OF WIND MOVEMENT
An experiment devised to test this factor was set up as follows:
A nursery flat about 24 inches square was filled with vermiculite. Pots
of plants, each about the same size, were placed in the vermiculite
spaced about as shown in Figure 1. The table under the flat was
marked A, B, C and D. The direction of the wind movement was at
side B. In the tests, larvae were placed in the central area, shown by a
small circle and allowed to go to the plants, or to leave the flat.
Ten different larvae were used for each experiment, and these were
used a total of ten times each. Thus, each experiment involved 100
trials.
The wind direction relative to- the position of the plants was
changed by turning the flat by 90° after each 100 trials, as shown by
the illustration (Fig. 1). The first position tested was with mustard
toward the dirction from which the breeze came ( side B ) . The results
of the preferential selection by the larvae are shown on Table 1.
Kale was selected by 19%- of the larvae, mustards by 56%, radish
1 Aided by a grant from the National Science Foundation, Washington, D.C.
2The authors wish to express their great appreciation to Dr. W. S. Stewart, Director, Los Angeles
State and County Arboretum, for great cooperation in making this work possible.
51
52
HOVANITZ AND CHANG
/. Res. Lepid.
TABLE 1. The effect of wind movement on the choice of food
plants by larvae of Pieris rapae
Kale
Mustard
Radish
Nasturtium
Isomeris
None
Total
no. of
trials
A 19
B56
C18
D3
D 1
3
100
90°
D20
A 20
B52
C4
C2
2
100
90°
C43
D25
A 26
B2
BO
4
100
90°
B53
C 15
D13
A8
A5
6
100
Ten 14.1 mm to 15.7 mm kale-bred larvae of Pieris rapae were used per test
with each test ten times at each position. The parents of the larva were collected
at western Orange County in a cabbage field.
by 18%, nasturtium by 3% and Isomeris by 1%. Larvae which left
the flat without selection amounted to the additional 3%.
With the flat turned 90” so that the radish was now in position B
( in the direction of the wind ) , the selection results were very different.
Kale remained relatively unchanged at 20%, mustard dropped from
(i);5i~6i, 1962
THREE FACTORS
53
56% to 20%, radish rose from 18% to 52% at position B, while the
remainder were unchanged.
With a further 90° turn in the direction of the arrows (Fig. 1),
the radish plant was now opposite the direction of the wind, and
nasurtium-Isomeris was toward the wind. Kale at position C to the
right side now rose from 20% to 43%, mustard at position D (away
from the wind) rose slightly and v/as now 25%, radish at the left side
was now 26% and nasturtium-Isomeris (toward the wind) was still
relatively unchanged at 2 % .
The final 90° turn placed kale in the position toward the wind.
Here, 53% of the larvae went to kale, 15% to mustard (right side),
13% to radish (opposite side), and 13% to nasturtium-Isomeris
(left side) .
CONCLUSIONS:
( 1 ) Of the four sides of the flat which could face the direction
of the wind, the plants on that side were favored in three of the four
arrangements, namely, mustard, radish and kale. Only when the
nasturtium-Isomeris combination v/as toward the wind was the fre-
quency in position B not increased. In fact, at that time, it was
decreased.
(2) These tests, of course, cannot indicate which of the three
favored plants are relatively more favored since they each received
about the same percentage of selection when toward the wind. However,
when away from the wind, radish received fewer selections than either
kale or mustard.
(3) The Isomeris-nasturtium combination received its highest
selection when in position A (left side). In this position also, more
larvae left the flat without a plant selection than in any other position.
(4) The higher selection of kale (43%) while at position C
(right side), than radish (26%) while at position A (left side) seems
to indicate a preference for kale rather than for radish.
( 5 ) The lowest selection for the nasturtium-Isomeris combination
was when it was in position B (toward the wind). This would seem
to indicate that these plants (and Isomeris especially) repel rather
than attract larvae of Pieris rapae.
(6) The data indicate that the direction of wind movement is of
great importance in larval selection of food plant, since in all cases
where the plant has a positive effect on selection, that position in the
direction of the wind has by far the greatest selective influence. It
is apparent that the reverse is also true, that if a plant is repellent,
the effect is greatest when the wind comes from that direction.
[NOTE: The differences between the figures indicated are so great that
the element of chance being involved is negligible. A chi-square test on the
first trial gives a probability of less than one in a million that the differences are
due to chance alone.]
54
HOVANITZ AND CHANG
/. Res. Lepid.
EFFECT OF LARVAL SIZE ON PLANT SELECTION
Tests for the determination of influence of larval size on plant
selection were made in the same manner as indicated previously, except
that all tests were carried out in a room with no wind or breezes.
Nevertheless, the flat was turned 90° three times for each test. This
applies to all subsequent tests for the sake of safety. Two . series of
tests were made, using larvae of different origin.
The first series of tests (Table 2) was carried out with the use
of kale-bred larvae. Larvae of different ages and sizes (size is roughly
proportional to age) were separated and measured. Four larvae were
isolated, these being the following sizes:: 8.4, 14.7, 19.0 and 21.0 mm
in length. The tests were carried out by five trials of a given larva
at each position with respect to the direction of the wind (Fig. 1),
giving a total of 20 trials for each size larva.
TABLE 2. Choice of five kinds of plants by different sized larvae
of Pieris rapae ( Series 1 )
Size of Total
larvae The number of times and percentage larva goes to Number
(mm)
Kale
Mustard
Radish
Nasturtium
Isomeris
None
of trials
21.00
16- — 80%
4—20%
0— 0
0—0
0—0
0—0
20
19.00
13—65%
3—15%
3—15%
1—5%
0—0
0—0
20
14.70
11—55%
4—20%
4—20%
1—5%
0—0
0—0
20
8.40
10—50%
5—25%
5—25%
0—0
0—0
0—0
20
One single kale-bred larva was used for each size; tested each larva five times
at each position.
The 8.4 mm larva selected kale 50%- of the time, radish 25% of the
time and mustard 25% of the time.
The 14.7 mm larva selected kale 55%, mustard 20%,. radish 20%
and nasturtium 5 % of the time.
The 19.0 mm larva selected kale 65%, mustard 15%, radish
1 5 % and nasturtium 5 % of the time.
The 21.0 mm larva selected kale 80% and mustard 20% of the
time and no others.
The probability, as caculated by chi-square, that the differences
as indicated might be due to chance are less than 1/million, 1/100,000,
1/500, 1/500 respectively for each experiment.
CONCLUSIONS:
( 1 ) The increase in selection of kale for each experiment is
correlated with increase in size of the larvae ( Fig. 2 ) .
(2) Kale is the plant selected over all the others in this experi-\
ment. It should be noted that the larvae had been bred on kale prior
to the experiment, and previous generations had been bred on cabbage.
(3) Not one larva went to Isomeris'. This is interesting in view
of the observation in the preceding experiments that Isomeris was
(i);5i-6i, 1962
THREE FACTORS
55
2. The relation between size (and age) of larvae and the percentage of the
time kale was selected. The size of the larvae is directly correlated with an
increase in selection of kale.
repellent to Pieris rapae larvae.
(4) The older or larger the larvae, the greater the selection of
kale over all other plants ( Tig. 2 ) . The relationship is not a straight-
line, but rather a geometric increase. This indicates that there is an
increase in the critical powers of food perception in the older or
larger larvae.
In the Series No. 2 tests (Table 3), larvae were used which had
previously been grown on nasturtium. They were obtained from
Ventura County on nasturtium and were continued in the laboratory on
that plant.
Two larvae 22.4 mm in length, and four larvae 16.5 mm in length
were used in this experiment. Otherwise, the trials were managed
in the same way as in previous experiments with the exception that
the total number of trials was 80 for large larvae and I6O for the
56
HOVANITZ AND CHANG
/. Res. Lepid.
smaller larvae.
The selection by these larvae bred on nasturtium (Table 3)' was
radically different from the selection by larvae bred on kale ( Table 2 ) .
The selections of kale by the small larvae bred on nasturtium were
15% as compared with 55% for a comparable size bred on kale. The
selections of kale by the large larvae bred on nasturtium were 22%
compared with 80% for a comparable size bred on kale. The greatest
shift in food plant selections for the nasturtium-bred small larvae
was 49% to mustard as compared with 25%, 9% to nasturtium as
compared with 0%, and 16% with no selection as compared with 0%.
For the large larvae, comparable data were 34% to mustard
as compared with 20%, 33% to nasturtium as compared
with 0%, and 9% with no selection as compared with 0%.
CONCLUSIONS:
( 1 ) The shifts in food-plant-selection by the larvae grown on
different plants are highly significant and indicate a direct relationship
between the feeding habits of the larvae, and their selection when
given a free choice.
(2) The direct relationship between the size (and age) of the
larvae and their selection of the plant previously eaten is clearly
indicated.
(3) A plant previously of nearly negligible selection may be the
most highly selected if the larvae have been grown on that plant. For
example, larvae selected nasturtium over all others (33%) after being
bred on that plant (Table 3), whereas larvae bred previously on kale
had almost no selectivity for that plant.
INFLUENCE OF PREVIOUS FOOD PLANTS OF LARVAE
Additional experiments to test the influence of previous exposure
to larval food plants are reported on here.
The first comparative test involves two strains of Pieris rapae, one
from a population growing in the wild on black mustard and the other
growing on cabbage. These strains were kept in the laboratory for
several generations; the mustard strain was bred for over six genera-
tions on mustard in the laboratory and the kale strain was bred on kale
in the laboratory for over ten generations.
Larvae were selected for these experiments which had a size of
14 to 16 mm. Using twenty-five larvae of each strain, 600 tests of
selection were made (Table 4) : the selected plants were mustard,
kale, nasturtium, Isomeris and Cleome.
The only significant differences detected in the tests between the
two strains were the relative selections made of mustard and kale.
Larvae from the mustard-bred strain selected mustard 61% of the time
compared with the kale-bred strains of larvae of 24% on that plant.
On the other hand, kale was selected by the mustard-bred larvae 20%
TABLE 3. Choice of fi¥e kinds of plants by different sized larvae
of Pieris rapae ( Series 2 )
57
1^62
THREE FACTORS
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The nastuf tram-bred larvae were from Ventura County, collected in a nasturtium
garden. The parents of the kale-bred larvae were collected io western Orange
County on cabbage; the larva are the first generation fed on kale.
58
HOVANITZ AND CHANG
/. Res. Lepid.
of. the time as compared with 59% by the kale-bred larvae. Testing
these differences by chi-square using the four-fold table indicates a
probability of less than 1 /million that the differences could be due to
chance alone. This may be calculated in another way, that is, by taking
364 + 354 = 718 for one value and 119 + 144 = 263 for the other.
The expectation that these differences might be due to chance alone,
that is, that the deviations 718 — 981, and 971 — 263 are not due to
2 2 ‘
chance is well over one in a million.
CONCLUSIONS:
( 1 ) The shift in selection of mustard or of kale according to
whether or not the larvae were from a kale or mustard strain is
indicated by the experiments. The data are highly significant at an
incredibly high level.
( 2 ) There is no significant change indicated in the selective level
of the other plants involved as to whether or not the larvae were from
the mustard or kale strain.
The second series of data on the problem of food plant selections
involves nasturtium-bred larvae from a nasturtium strain and kale-
bred larvae from a cabbage strain (Table 5). The experiments were
carried out as before, two larvae of each strain were used, the same
size for a total of forty trials each, of which ten were in each position
as indicated in Figure 1.
The differences between the plant selections are again highly
significant. The nasturtium -bred larvae as compared with the kale-
bred larvae preferred mustard (35% as compared with 22.5%) and
nasturtium (15% over 4%). The kale-bred larvae preferred kale
(49% over 12.5%) and radish (22.5% over 14.0%). Neither
selected Isomeris to any extent. A high proportion (22%) of the
nasturtium-bred larvae rejected all plants.
CONCLUSIONS:
( 1 ) As in previous experiments, there was a shift in selection by
the larvae according to the previous larval food plants.
( 2 ) This shift in selection was in the direction of the plant they
had previously fed on. For example, nasturtium-bred larvae selected
nasturtium better than did kale-bred larvae.
( 3 ) Even though nasturtium-bred larvae selected nasturtium
better than did kale-bred larvae, mustard was preferred over nasturtium
and over kale.
(4) The greater selection of mustard over kale bv nasturtium-
bred larvae is significant not only in this experiment but also in a
previously indicated test (Table 3). In both cases, kale is poorly
preferred as compared with mustard when the larvae were nasturtium-
bred.
(5) Radish is of lower selective influence when the larvae are
(i):5i-6i, 1962
THREE FACTORS
59
nasturtium-bred than when they are kale-bred. This was indicated
also in previous tests ( Tables 3 and 1 ) .
(6) A significantly higher percentage of rejects (22%) are
made by the nasturtium-bred larvae than by the kale-bred. This was
also indicated in previous tests (Table 3 as compared with 1, 2 or 4).
DISCUSSION
All three factors, investigated as to their relevance in influencing
larval choice of food, have been shown to be important in that regard.
These factors are:
( 1 ) Influence of air movemenr.
(2) Influence of age (size) of larvae.
(3) Influence of food previously eaten by larvae, or strain of
larvae.
In addition, by virtue of the laboratory set-up used in these
experiments, some further conclusions may be drawn which are not
completely in accord with some previous conclusions reached by others.
One of these further conclusions is with regard to the ability of
lepidopterous larvae to "recognize” food plants from a distance. In the
present experiments, the larvae were attracted to the plants from a
distance of over 120 mm without any difficulty and responded in most
cases immediately.
Chin (1950) reports that the larvae of the Colorado potato beetle
(Leptinotofsa) could not perceive its food plant from a distance farther
than 2mm. Gupta and Thorsteinson (I960) tested the olfactory
response of the larvae of the diamond backed moth [Plutella macul-
ipennis (Curt.)] toward mustard oil odors and found that 5 mm was
the maximum distance of response. Dethier (1959) in a study on
food plant distribution and density and larval dispersal as factors
affecting insect populations, states in relation to field observations that
"Larvae find new food-plants by chance alone.” It is presumed that
he had reference to the gross search for new food plants, not the
closer relationship. In the case of the experiments of Chin, and Gupta
and Thorsteinson, the screen test was used. In this, the larvae were
placed on a screen above the food plant or other attractive source. In
the present experiments, the larvae were below the food plants. This
difference could account for a fundamental difference when testing for
an odiferous substance heavier than air. Under natural conditions, our
observations have always indicated that larvae tend to climb upwards
rather than down so that tests involving simulated natural conditions
should be best performed with the attraction above rather than below
the larvae (Fig. 3). This point will be covered more fully in tests
to be described in later papers.
In a paper available to us after the tests here indicated were
performed, (Takata, 1959), there has been indicated a change in food
plant preference of larva when reared successively on different food
60
HOVANITZ AND CHANG
/. Res. Lepid.
3. Larva of Pieris rapae on edge of pot during one of the selection experiments
reaching out to black mustard plant three to four inches away. The larva
had already traveled several inches under directed impulse to get this far.
plants for several generations. These results complement ours, even
though the technique of testing used by Takata was different. He used
cabbage and radish with Pieris rapae crucivora larvae. In testing, he
used actual feeding experiments of the weight of leaves of the plants
eaten by the larvae.
(i):5i-6i, 1962
THREE FACTORS
61
SUMMARY
The evidence in this paper indicates ( 1 ) that larvae of Pieris rapae
are capable’ of detecting their preferred food plant from a distance of
at least 120 mm under the conditions of these experiments, (2) that
the direction of wind movement is very important to detection of food
plant, presumably through odor, (3) that the older and larger larvae
are much better at selecting the preferred food plant at least at a
distance and (4) that the influence of food previously eaten by the
larvae, or the strain of the larvae, directly influences their choice of
food in the direction of the previously eaten food.
LITERATURE CITED
CHIN, CHUN-TEH. 1950. Studies on the physiological relations between
the larvae of Leptinotarsa decemlineata Say and some solanaceous plants.
H. Veenman and Sons, Wageningen, Holland, pp. 1-88.
DETHIER, V. G. 1959. Food plant distribution and density and larval
dispersal as factors affecting insect populations. Canad. Entom. 91: 581-596.
GUPTA, P. D. AND A. J. THORSTEINSON. I960. Food plant relation-
ships of the diamond-backed moth (Plutella maculipennis (Curt.)). I.
Gustation and olfaction in relation to botanical specificity of the larva.
Ent. exp. and appl. 3 (I960) :24l-250.
TAKATA, N. 1959. Studies on the hobt preference of common cabbage
butterfly, Pieris rapae crucivora (Boisduval). VI. Change in the food
preference of larvae when reared successively by the definite food plant
for several generations (preliminary report). Jap. J. Ecol. 9:224-227.
1(1) :63-71, 1962
Journal of Research on the Lepidoptera
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Copyright 1962
THE GENERIC, SPECIFIC AND LOWER CATEGORY
NAMES OF THE NEARCTIC BUTTERFLIES
PART 1 — The Genus Pieris
PADDY McHENRY
1032 E. Santa Anita, Burbank, California
PREFACE
INTRODUCTION
The author has asked that I write a brief introduction to this,
the first of a series of compilations of generic, species and lower
category names of the Nearctic butterflies. The objects of the work are
(1) to give a complete list of all the names proposed for the butterflies of
the Nearctic region from the year 1738 until the date of publication
of the list, (2) to indicate the date and place where originally published,
(3) to indicate the types of the genera, species and lower category
names where known, (4) to indicate the type locality in the original
author’s words for types, except those for the generic names, and
(3) to indicate the location where types have been deposited where
indicated by the author.
The biological application of these names so indicated is not to
be attempted in this series. Instead, this effort is left to the individual
who makes a comprehensive study of the biology of the groups con-
cerned, such a person rarely having the time and inclination to make
the detailed and laborious studies of nomenclature as such. There
is urgent need for a work such as this even in a field as well civilized
as the Lepidoptera; in fact, this kind of work is so long overdo that
the civilized state is rapidly reverting to chaos. Each new volume
which appears from the press on this group of insects plows under well
established names with no reason given to justify such destruction.
A most valuable start in the direction of making reason out of chaos
was started by Francis Hemming in his TThe Generic Names of the
Holarctic Butterflies” published in 1934. This work is proceeding
also in the sam.e direction.
When the various parts of this work are completed, it is expected
to reissue the whole as a separate volume. Then, perhaps we can
prevail upon Mr. McHenry to go on to a similar effort with the moths,
William Hovanitz
63
64
McHENRY
/. Res. Leptd.
Each entry for the generic names includes the following; the
name, the author’s name, the most accurate determinable publication
date, the bibliographical reference (including the volume or subvolume
for the date indicated ) , the type species, the selection of the type species
and the supporting data to justify the type indicated.
Each entry for the specific names includes the following; the name
as originally given, the author’s name, the most accurate determinable
publication date, the bibliographical reference, data on type locality, sex,
season, etc., other spellings of the name and notations of any homonym.
The date of publication of a name is indicated in a manner that
shows its relative accuracy; for example, "mailing” and "release” dates
if known are given preference over "publication” dates if they are not
the same. All dates are given in English and normally in the sequence
of day, month and year. Where more than one date is indicated, the
more accurate one is given after the less accurate one. Dates ascertained
directly from a source whose purpose was the determination of time of
publication of a work are given without brackets; others, ascertained less
directly are given in brackets and explained in a footnote. Combinations
of dates with and without brackets may be given when necessary.
I have seen originals or photostats of all references except as noted
but have been unable to ascertain the contents and dates for the various
parts of Verity’s Rhopalocera Palaearctica.
The names of the genus Pieris are listed under the five species
indicated, since in this genus the separation does not involve great
difficulty. I have been aided in this regard by William Hovanitz, or
by reference to the original description. Should these be in error,
no harm is done since the total list, not the biological interpretation of
the categories to which the names applied, is the purpose of the listing.
Should anyone find names which have been inadvertently omitted
from this listing, a great service will be rendered to so notify the author
or the editor of this fact so that corrections may be made.
I take this opportunity to express my gratitude for the use of
facilities and to the staffs at the Allan Hancock Foundation Library of
the University of Southern California, and at the Los Angeles County
Museum Library & Entomological Department. A debt of gratitude is
tenderd to the photographic service sections of the U. S. Department of
Agriculture Library, Harvard College Library, New York Public Library,
University of California Library and Cornell University Library. An
extended exchange of correspondence with Mr. C. F. DosPassos has
been most helpful.
LIST OF GENERIC NAMES USED OR
AVAILABLE FOR PIERIS
ASCIA Scopoli.
Type; monuste L.
(i):63-7I, 19^2
NAMES Pt. 1
65
MANCIPIUM Hubner.
Type: hellica Hubner = hellica L. = helice L.
PIERIS Schrank.
Type: hrassicae of var. authors = brassicae L.
ANDROPODUM Hubner.
Type: brassicae Hubner = brassicae L.
GANORIS Dalman.
Type: brassicae L.
TACHYPTERA Berge
Type: brassicae L.
PONTIA Fabricius.
Type: daplidice Fab. = daplidice L.
SYNCHLOE Hubner.
Type: callidice Butler = callidice Esper = callidice Hubner
ASCIA SCOPOLI. 1777. Introd. Hist. Natur. : 434, no. 175. Includes
among others no. 80 (="P. D. [C.] monuste” of Linnaeus, 1767)
which in turn is Papilio D. C. monuste Linnaeus, 1764.
Type Papilio D\anaus\ C[andidus]. monuste Linnaeus. 1764. Museum
Ludovicae Ulricae ([I]) : 237, no. 56.
Type Selection. Scudder. 1872 [in or pre-June] L Syst. Rev. Some Amer.
Butt., p. 40. Says of Ascia: "Type Papilio monuste Linn.”
MANCIPIUM HUBNER. [1 Jan. 1807] - [19 Dec. 1807]L Samrn. Exot.
Schmett, 1: pi. [141], ff. 1-4. Gives "Mancipium vorax hellica”. As
used by Hubner, hellica is considered to be hellica Linnaeus, 1767
which in turn is helice Linnaeus, 1764.
Type. Papilio D[anaus]. C[andidus]. helice Linnaeus. 1764. Museum
Ludovicae Ulricae ([I]) : p. 243, no. 62.
Type Selection. Hubner. As above. Gives only hellica, at the time,
which becomes the type.
PIERIS SCHRANK. 1801. Fauna Boica 2(1): 152, no. 198; 2(1):
160-170, nos. 1283-1296. Includes Pieris brassicae which he says is
Papilio brassicae of Goeze, Borkhausen, Brahm, and Esper. Of these
authors, brassicae is considered to be hrassicae Linnaeus, 1758.
Type. P\ap)ilio\ D\anaus\ [Candidas] hrasicae Linnaeus. 1758. Syst.
Nat. 10th Ed. 467-468, no. 58.
Type Selection. Latreille. 1810. Consid. Gen. Anim. Crust. Arach. Ins.
p. 440. Says: "Pieride. Pontia brassicae, Pab.”. As used by Latreille,
brassicae Fabricius !s considered to be brassicae Linnaeus, 1758.
ANDROPODUM HUBNER. 1822. [post 22 Sept.]." Syst.-Alph. Verz. pp.
2-5, 7-9. Includes "Brassicae L, 401-403. Andropodum vorax.”
Type. P\apilio\ D[anaus]. [Candidas] hrassicae Linnaeus. 1758. Syst.
Nat. 10th. Ed. 1:467-468, no. 58.
Type Selection. Hemming. Sept. 1933. Entomologist: 66(844) : 199.
Says of Andropodum: "Type =: Andropodum hrassicae Linn.”
GANORIS DALMAN. 1816. Kongl. Vetenskaps Academiens Handlingar
1816(1) : 61-62, no. 8; 1816(1) : 86-90, nos. 1-10; tablet 1, no. 8;
tablet 2, no. 8. Includes G. brassicae in which he cites Linnaeus
Fauna Sv., pp. [269] -[2701, no. [10351 (this is hrassicae Linnaeus,
1761, which is hrassicae Linnaeus, 1758).
Type. Plapilio]. D[anaus]. [Candidas] Brassicae Linnaeus. 1758. Syst.
Nat. 10th Ed. 1:467-468, no. 58.
Type Selection. Dalman. As above. Says of Ganoris: "Generis Typus: Pap-
hrassicae”.
66
McHENRY
/. Res. Lepid.
TACHYPTERA BERGE. 1842. Schmetteclingsbuch, p. 19, pp. 92-105.
Not seen, given as per Hemming, 1934.
Type. P[apilid\, .Dlanaus\ [Candidus] brassicae Linnaeus. 1758. Syst.
Nat. 10th. Ed. 1:467-468, no. 58.
Type Selection. Hemming. Feb. 1934. Entomologist: 67(849) :38. Says
of Tachyptera: "Type Papilio brassicae Linn., 1758.”
PONTIA FABRICIUS. 1807. In Illiger. Magazin fur Insektenkunde. 6:
283, no. 23. Includes: ’Tap . , . Daplidice”. As used by Fabricius
daplidice is considered to be daplidice Linnaeus, 1758.
Type. P[apilio]. D[anaus]. [Candidus] daplidice Linnaeus. 1758. Syst.
Nat. 10th Ed. 1:468, No. 2.
Type Selection. Curtis. 1824. Brit. Entom. 1: pi. 48. Not seen,
given as per Hemming, 1934.
SYNCHLOE HUBNER. 1818 [post 22 Dec.]^. Zutr. Samm. Exot. Schmett.
1st. 100 (text in 1): p. 26, under no. 76. (Synchloe autodice).
Includes "S. callidice”. Refers to his earlier publication of this
butterfly.
Type. Papilio callidice Hifbner. [24 Dec. 1799] - [13 Apr. 1800]^. Samm.
Europ. Schmett. (Papiliones) : plate 81, figs. 408-409. Page 63, no. 5
and plate 108, figs. 551-552 were published later.
Type Selection. Butler. 12 Sept. 1870. Cistula Ent. 1(3):51. Says of
Synchloe: "Type S. callidice Esper”. As used by Butler callidice Esper
is considered to be callidice Hubner [24 Dec. 1799] - [13 Apr. 1800].
LIST OF SPECIES AND LOWER CATEGORY NAMES
USED OR AVAILABLE FOR PIERIS
1. P. (PONTIA) BECKERII EDWARDS.
heckerii Edwards.
gzmderi Ingham.
pseudochloridice McDimnough.
2. P. (PONTIA) SISYMBRII BOISDUVAL.
elivata ( B. & B. ) .
jlava Edwards.
flavitincta Comstock, J. A.
sisymhrii Boisduval.
transversa {B. 8iP>.)
3. P. (PONTIA) PROTODICE BOISDUVAL & LECONTE
calyce Edwards. .
nastuftii Edwards ( Bdv. Ms. ) .
nehoni Edwards.
occidentalis Reakirt.
protodice Boisduval & LeConte.
vernalis Edwards.
4. P. (PIERIS) NAPI (LINNAEUS).
acadica Edwards.
aestiva Edwards.
arctica Verity.
borealis Grote.
casta Kirby.
i(i):6}-7T, 1962
NAMES — Pt. 1
67
castoria Reakirt.
cottlei Gunder.
cruciferamm Boisduval.
flava Edwards.
flava Edwards.
ffigida Scudder.
hulda Edwards.
hyemalis Edwards.
iberides Boisduval.
macdunnoughii Remington.
marginalis Scudder.
micro striata Comstock, J. A.
mogollon Burdick.
napi (Linnaeus).
nasturtii Boisduval.
ochsnheimeri Staudinger.
oleracea Harris.
pallida Scudder.
pallidissima B. & McD.
pseudobryoniae Verity.
pseudoleracea Verity.
pseudonapi B. & McD.
resedae Boisduval.
venosa Scudder.
virginiensis Edwards. 1870.
virginiensis Edwards. 1881.
5. P. (PIERIS) RAPAE (LINNAEUS).
aestivus Verity.
immaculata De Selys-Longchamps.
immaculata Cockerell.
immaculata Skinner & Aaron.
metra Stephens.
novangliae Scudder.
rapae ( Linnaeus ) .
yreka Reakirt.
1. PIERIS (PONTIA) BECKERII EDWARDS.
heckerii Pieris Edwards. [Sept. 1871]*^. Butt. N. Amer. 1(8) : [28a] -
[29]; plate [81], figs. 4-7. 'cf described. "Virginia City, Nevada,
April 1870.” "Four individuals”. McDunnough, 1 Dec. 1928, spells
heckeri.
gunderi, Ascia heckerii Ingham. 14 Apr. 1933. Pan-Pacific Ent. 9(2) :75.
Holotype ? : "Bouquet Canyon, Los Angeles County, Calif”. "Bred
from larva”, "emergence date June 21, 1932”. Type deposited: Calif.
Acad. Sci., San Francisco, Calif.
pseudochloridice, Pieris heckeri McDunnough. 1 Dec. 1928. Can. Ent.
60(11) : 266-267. Holotype d* : "Oliver, B[ritish]. C[olumbia]., April 24
. . . No. 2861 in the Can. Nat. Coll., Ottawa”. Allotype ? : "Oliver,
68
McHENRY
/. Res. Lepid.
B.C., April 22”. Paratypes — I d*: "Hedley, B. C., May 15”. 1 9:
"Osoyoos, B.C.” 1 + : "Osoyoos, B.C.”
2. PIERIS (PONTIA) SISYMBRII BOISDUVAL.
elivata, Ascia sisymhni Barnes & Benjamin. 8 Dec. 1926. Bull. Sou. Calif.
Acad. Sci. 25(3) :88, no. 33a. "Type locality: Glenwood
Springs, Colo.”. "Number and sexes of types: Holotype d* May 1895,
Allotype ? May 1895, 7 d» 14 9 Paratypes, various dates April to
June”.
jlava, Pieris sisymhri Edwards. Apr. 1883. Butt. N. Amer. 2(11): [67];
plate [151. fig 5. ? described. No locality, series data nor dates given.
flavitincta, Pieris sisymhrii Comstock, J. A. 20 Feb. 1924. Bull. Sou. Calif.
Acad. Sci. 23(1) : 19-20; plate 7, fig. 9. 9 described. "The example
before us was captured . . . April 30th, 1911 at Cranbrook, British
Columbia”. Intends to propose new name for jlava Edwards.
sisymhrii, Pieris Boisduval. 1852 [25 Feb. - 22 Dec. Ann. Soc. Ent.
France. 2nd Ser. 10 (2) :284, no. 8. 9 described. [California]. No series
data nor dates given. Edwards, Apr. 1883, spells sisymhri.
transversa, Ascia sisymhrii Barnes & Benjamin. 8 Dec. 1926. Bull. Sou.
Calif. Acad. Sci. 25(3): 88, under no. 33. "Type localities: Paradise,
Cochise Co., Ariz.; Redington, Ariz.”. "Number and sexes of types:
Holotype cf , Allotype 9,1 9 Paratype, all March; 1 cf Paratype, no
date”.
3. PIERIS (PONTIA) PROTODICE BOISDUVAL & LECONTE.
calyce, Pieris Edwards. Nov. 1870. Trans. Amer. Ent. Soc. 3(?) sign. 25:
189, no. 1 (Nomen nudum); 189. d* described. "From Nevada; in the
collection of Henry Edwards”. No series data nor dates given.
nasturtii, Pieris Edwards (Boisduval Ms. name). [9 May 1864]°. Proc. Ent.
Soc. Phila. 2(4) : 501, no. 1 (Nomen nudum); 501. d* & 9 described.
"San Francisco; from Dr. Behr, who informs me that it is common in
some localities in the vicinity of that city”. No series data nor dates
given.
nelsoni, Pieris Edwards. Apr. 1883 Butt. N. Amer. 2(11): [71] - [72];
plate [15], figs. 6-7. "1 cf ... St. Michael’s Alaska, June, 1881”.
occidentalis, Pieris Reakirt. June 1866. Proc. Ent. Soc. Phila. 6 (pp.
121-152) : 133-134. cf & 9 described. "Hab. - Rocky Mountains,
Colorado Territory, California, (Coll. Tryon Reakirt)”. No series data
nor dates given.
protodice, P[ieris]. Boisduval & LeConte. 1829. Hist. Gen. Icon. Lepid-
Chen. I’Amer. Sept. 1 (5) :45-46. Plate 17, figs. 1-3 published later. A
& 9 described. "Elle parait au printemps et a la fin de juin aux
environs de New-York. Nous en avons vu un individu pris dans le
Connecticut”.
vernalis. Pieris Edwards. [9 May 1864]®. Proc. Ent. Soc. Phila. 2(4) : 501,
no. 2 (Nomen nudum); 2 (4) :501-502. cf & 9 described. "In the
collection of Mr. Geo. Newman and Mr. Wilt are several specimens,
taken, as I am informed, at Red Bank, New Jersey, in the month of
May”. "There is also in the Society’s collection a pair from the Rocky
Mountains, that appear to be identical with it”.
4. PIERIS (PIERIS) NAPI (LINNAEUS). _ .
acadica, Pieris napi Edwards. June 1881. Papilio l(6):86-88 (in pt.),
sub no. 1; p. 98, no. 1 of no. 3; p. 99; plate 3, figs. 10-11. It appears
that one pair was from Southern Newfoundland and was taken in the
last week in July, it also appears another set ( 1 cf 2 9 ) had emerged
from pupae in August and was from the same locality.
aestiva, Pieris napi oleracea Edwards. June 1881. Papilio l(6):86-88 (in
pt.. Nomen nudum), sub no. 1; l(6):89-95 (in pt.)
sub no. 3; l(6):95-98 (in pt.), sub no. 4; 1(6):98, no. 3 of no. 3;
i( 1962
NAMES — Pt. 1
69
1(6):99; plate 3, figs. 15-16. cf & described. "NewYork” (see
figs, explanation, p, 99). "summer brood”. No series data given.
arcUca, Pieris napi frigida Verity. 1905-1911. Rhopal. Palaearctica : pp.
333-334 (in pt., cites figs 32-33 on plate XXXII and figs. 16-17 on
plate LXVII as P. n. f. arctica)\ plate XXXII, figs. 32-33; plate
XXXII explanation page, figs. 32-33 (gives as P. n. frigida); plate
LXVII, figs. 16-17; plate LXVII explanation page, figs. 16-17 (gives as
P. n. arctica); p. xxviii (cites figs. 31-33 & 36-37 on plate XXXII
and figs. 16-17 on plate LXVII, gives as P. n. /. arctica). "31 . . . cf
(ile d’Yesso, Japon)”. "32 . . . c? , (Norvege sept.)”. "33 . . . d*
(Nulato, Alaska)”. "36 ... $ (Nulato, Alaska)”. "37 ... 9
.(Finmark, Scandinavie) ”. ”16 , 9 (Saltdalen, Norvege sept.) ”. "17
. . . ? (Laponie)”. Figs. 31, 33, 36: [coll, de Joannis]. Fig. 32: [coll.
Stefanelli]. Fig. 37: [coll. Obth.] Fig. 16: le coll. Mourray]. Fig. 17: [e
coll. Leach]. No series data nor dates given.
borealis, Ganoris oleracea Grote. Nov. 1873. Bull. Buffalo Soc. Nat. Sci.
1(4) sign. 24:185. "months of June and July, on the Island of Anti-
costi”. No series data nor sex given.
casta, Pontia Kirby, W. 1837. In J. Richardson. Fauna Boreali-America
(4) :288, no. 1; plate 3, fig. 1. "Three specimens taken in Lat. 65°”.
No sex given.
castoria, Pieris Reakirt. [11 June 1866] - [13 Feb. 1867]*^. Proc. Acad. Nat.
Sci. Phila. [18] (3) :238, no. 2. d* described. "Hab. - California. Coll.
Tryon Reakirt”. No series data nor dates given.
cottlei, Pieris napi castoria Gunder. 6 July 1925. Ent. ^ews 36(7) :197-
198, no. 9; plate 5, fig. 9; Holotype d : "(author’s Coll.), Anderson
Springs, Lake County, California, May 5, 1919”.
crucifer arum, Pieris Boisduval. 1836 [in or post Apr.]^°. Hist. Nat. Ins.
Spec. Gen Lepid. 1 (text in 1) :519, no. 119. d* described, "dans les
provinces septentrionale des Etats-Unis”. No series data nor dates given.
flava, Pieris napi pallida Edwards. June 1881. Papilio 1(6) :89”95 (in pt.,
unnamed), sub no. 3; 98, sub no, 2 of no. 3. "Washington Terr[itory].”.
"One of these [a $ ] is yellow on the upper side”. No date given.
flava, Pieris napi venosa Edwards. June 1881. Papilio 1(6) :88-89 (in pt,,
unnamed), sub no. 2; 98, sub no. 1 of no. 2. Fig. 7 of plate 2
(although not so named in plate explanation, p. 99) appears to be
flava. "A large percentage of female Venosa are yellow on the upper
side”. No locality, series data nor dates given.
frigida, Pieris Scudder. Sept. 1861. Proc. Boston Soc. Nat. Hist. (8(?)
sign. 12:181-182. "Two . . . males . . . two females”, "on Caribou
Island, Straits of Belle Isle”. No dates given.
hulda, Pieris Edwards. Sept. 1869. Trans. Amer. Ent. Soc. 2(?) sign.
48:370. "From Kodiak, 1 o* . Coll, of Henry Edwards”, No date given.
hyemalis, Pieris napi oleracea Edwards. June 1881. Papilio l(6):88-89
(in pt.. Nomen nudum), sub no. 2; 89-95 (in pt.), sub no. 3; 95-98
(in pt.), sub no. 4; 99; plate 2, fig. 8. d* &■ $ described. Directly and
indirealy cites material from New Hampshire, Massachusetts, Lake
Superior area and Mt. Hood, Oregon. No series data nor dates given.
iherides, Pieris Boisduval. [1869, pre 1 Nov.]^’. Ann. Soc. Ent. Belgique 12
(in 1):39, no. 9. d* & ? described. [California]. No series data nor
dates given.
macdunnoughii, Pieris napi Remington. 17 Sept. 1954. The Lepid News
8(3-4) :75. A new name for Pieris napi pseudonapi Barnes & McDun-
nough, 5 Dec. 1916. See pseudonapi for series data.
marginalis, Pieris Scudder. Stpt, I86I. Proc. Boston Soc. Nat. Hist. 8(?)
sign. 12:183. "(Id* ,1 9 ) which are in the Museum of Comparative
Zoology”. The male . . . from Gulf of Georgia, and the female from
Crescent City, California”. No dates given.
70
McHenry
/. Re$. Lepid.
mkrostfiata, Pieris napi Comstock, J. A. 12 Sept. 1924. Bull. Sou. Calif.
Acad. Sci. 23(4) :125. Holotype f:- "Eldredge, Sonoma County, Cali-
fornia. March 13, 1911”. Allotype [•$■]: “same locality and on the
same date”. 'Taratype; one male - same locality and date”.
mogollon, Pieris napi Burdick. 28 Sept. 1942. Can, Ent. 74 (8) :154-155.
Holotype : "Mogollon Range, Catron Co., New Mexico, 5»7-40. No.
5222 in the Can . . . Nat . , . Coll . . . , Ottawa”. Allotype $ : "same
data”. "Paratypes. 2 ^ ,same data; 3 cf , Sierra Blanca, Lincoln Co.,
N. M., 5-25-40; all in the Can , . . Nat . . . Coll . . . , Ottawa. 1 d* ,
Mogollon Range, Catron Co., N. M., 5-7-40 and 1 $ , Sierra Blanca
Range, Lincoln Co., N. M., 5-20-40; in the U , . . S . . .^at . . . Mus . .
Fifty-six paratypes from the above locations in the collection of the
author, from which other museums will be supplied”.
napi, P[apilio]. Dlanausl. [Candidus] Linnaeus. 1758. Syst. Nat. 10th. Ed.
1 ;468, no, 60. No locality, series data, sex nor dates given.
Edwards, June 1881, spells napae.
nasturtii, Pieris Boisduval. [1869, pre 1 Nov.]^L Ann. Soc. Ent. Belgique
(12) (in 1) :38-39, no. 7. described, "dans les champs decouverts au
pied de la Juba”. "Nous o’avons vu que des males”. No series data
dates given. A homonym of Pieris nasturtii Edwards, [9 May 1864].
ochsenheimeri, Pieris Staudioger. End Apr. 1886. Ent. Zeitung Stettin
47 (4-6) : 199-200. d & 9 described, "wo sie die beiden Haberhauer in
Anzahl Ende Juni bei Namagao, jedenfalls hoch in den Gebirgen ge
fangen batten. Auch Maurer sandte mir in demselben Jahre ein im
Alai-Gebirge (sudlich von Margelan) gefangeoes + ein”. No series
data given.
oleracea, Pontia Har^fs. 10 July 1829. The New England Farmer & Horti-
cultural Jour. 7(51) :402. "Habitat ... in New Hampshire, and Massa-
chusetts”. No series data nor sex given, "two broods in a season”:
[May- June - August].
pallida, Pieris Scudder. Sept. 1861. Proc. Boston Soc. Nat. Hist. 8 (?) sign.
12:183. "five specimens (3 d* » 2 9.), which are in the Museum of
Comparative Zoology”. "Gulf of Georgia”. No dates given.
pallidissima, Pieris napi Barnes & McDunnough. 5 Dec. 1916. Contrib.
Nat. Hist. Lepid N. Amer. 3(2) :59; 142; plate 6, figs. 4-5 & 10.
"second generation (July, August)”, "type cf and $ from Provo,
Utah”. No series data given.
pseudobryoniae, Pieris napi frigida Verity. 1905-1911. Rhopal.- Palaearctica :
p. 146 (cites figs. 46-37 on plate XXXII, gives as P. n. f. pseudo-
bryoniae); plate XXXII, figs. 36-37; plate XXXII explanation page,
figs. 36-37 (gives as P. n. frigida); p. xxviii (cites only fig. 37, gives
as P. n. f. arctica pseudobryoniae) . "36 ... 9 (Nulato, Alaska) [coll,
de Joannis]”. "37 . . (Finmark, Scandiriavie) [coll. Obth.]”. No
series data nor dates given.
pseudoleracea, Pieris napi frigida Verity. 1905-1911. Rhopal. Palaearaica:
p. 146; plate XXII, fig. 38; plate XXXII explanation page, fig. 38;
p. xxviii. "38 . . . d' (Labrador) [e- coll. Boisduval in coll, obth.]”.
No series data nor dates given.
pseudonapi, Pieris napi Barnes & McDunnough. 5 Dec. 1916. Contrib.
Nat. Hist. Lepid. N. Amer. 3(2) :57; 142; plate 6, figs. 1-3. "type ^
and 9 taken at Silverton, Colo ... in the last week of July”. No series
data given. A. homonym of Pieris melete pseudonapi Verity, 1905-1911.
resedae, Pieris Boisduval. [1869, pre 1 Nov.]^ ’ Ann. Soc. Ent. Belgique
12 (in 1) :38-39, no. 8. 9 described. "Nous ne connaissons cette espece
que par un seul individu 'femelle pris . . . sur les bords du Sacramento”.
No date given.
venosa, Pieris Scudder. Sept. 1861. Proc. Boston Soc. Nat. Hist. 8(?)
i(i):6}-ji, 1^62
NAMES — Pt. 1
71
sign. 12:182-183. "twenty specimens (5 cf , 15 9 ), brought to the
Museum of Comparative Zoology, from San Mateo and Mendocino city,
California”. No dates given.
vkginiensis, Pieris Edwards. Jan. 1870. Trans. Amer. Ent. Soc. 3(?) sign.
2:10, no. 5 (Nomen nudum); 13-14. cf & $ described. "Not uncom-
mon in the Kanawha district [West Virginia] in the month of May, and
there replacing Oleracea. I have received from Mr. Saunders occasionally
specimens taken by him at London, Canada”. No series data given.
virginiensis, Pieris napi oleracea hyemalis Edwards. June 1881. Papilio
1 (6) :95-98 (in pt.), sub no. 4; 98, under no. 2 of no. 2. " . . . Virgin-
iensis . . . has become a true species, although unquestionably, in a higher
latitude, it appears as an occasional aberration only of Oleracea". Speci-
men data is obscure. A homonym of virginiensis Edwards, Jan. 1870.
5. PIERIS (PIERIS) RAPAE (LINNAEUS).
aestivus, Pieris rapae Verity. 16 May 1913. Jour. Linn. Soc. (London),
Zool. 32 (215) :177-178. "summer brood”. No locality nor series data
given.
immaculatag Pieris rapae De Selys-Longchamps. 1857. Ann. Soc. Ent.
Belgique 1 (in 1) :5, under no. 5. [Belgique]. No series data, sex nor
dates given.
immaculata, Pieris rapae Cockerell. Apr. 1889. Entomologist: 22(311) :99.
No locality, series data, sex nor dates given. Cites p. I6l in Newman’s
Brit. Butt. A homonym of Pieris napi immaculata De Selys-Longchamps,
1857.
immaculata, Pieris rapae Skinner & Aaron. 2 July 1889. Can. Ent. 21(7) :
128-129. "five specimens in the collection of Am. Ent. Soc., Dr. Skin-
ner and E. M. Aaron”. No sex nor dates given. A homonym of Pieris
rapae immaculata De Selys-Longchamps, 1857 and Cockerell, Apr. 1889.
metra, Pontia Stephens. 1 Aug. 1827. Illust. Brit. Ent. 1(?) sign, d: 19-20,
no. 4:146-147 published later, cf & ? described. "The insect occurs
early in April and a second time towards the end of June. I obtained
specimens of the first brood at Hertford; and of the second I captured
some this season, at Ripley at the latter period”.
novangliae, Ganoris rapae Scudder. Apr. 1872. Can. Ent. 4(4) :79. "Two
of my specimens were taken in June and July, 1869 — - one ... in
Shelbourne, N[ew]. H[ampshire] . . . the other ... in Waterville,
M[aine]., and all are males”.
rapae, P[apilio]. D\anaus\ [Candidusi Linnaeus. 1758. Syst. Nat. 10th. Ed.
l(in 1):468, no. 59. No locality, series data, sex nor dates given,
yreka, Pieris Reakirt. [11 June 1866] - [13 Feb. 1867]'^. Proc. Acad. Nat.
Sci. Phila. [18](3):238, no. 1. cf & 9 described. "Hab. - California.
Coll. Tryon Reakirt”. No series data nor dates given.
lAmer. Nat. 6: 3S4-559. Work is reviewed in June 1872 number.
^Hemming, 1937. Hubner 1:327-437.
^The work title page signature date qualified by preface signature date on page vi.
•^Title page signature date qualified by preface signature on page 6.
■^Hemming. 1937. Hubner. 1:146-324.
^Hemming. 1931. Proc. Ent. Soc. London. Ser. A. 6: 42-45. Gives actual dates of publication for
the parts of Edwards’s Butt. N. Amer., (Ser. I).
7Tome 10 title page signature date qualified by the meeting date of 25 Feb. (p. 275) and the
evidence indicating that the tome to page 489 was published before the meeting of 22 Dec.
(p. Ixxxii in Bulletin).
SProc. Ent. Soc. Phila. 1864. 3: p. 695. Notes that No. 4 of Vol. 2 of Proc. Ent. Soc. Phila.
was received at 9 May 1864 meeting.
9lndex Scient. Cont. Jour. & Proc. Acad. Nat. Sci. Phila., 1812-1912: pp. xii-xiii. Gives dates of
receipt for various parts of the early volume of the Proceedings.
1 OTome I title page signature date qualified by preface date (p. xii).
' 1 Trans. Ent. Soc. London 1869; p. xix (in Proc.). Notes receipt of Tomes 1-12 of the Ann.
Soc. Ent. Belg. at 1 Nov. meeting. Tome 12 title page signature date: 1868-1869. I assume the
date is fixed on the publication of Tome 12, I have no data for the separate except that its title
page signature date is 1869.
Journal of Research on the Lepidoptera 1(1) :73-83, 1962
1140 W. Orange Grove Ave., Arcadia, California, U,S.A.
Copyright 1962
THE DISTRIBUTION OF THE SPECIES
OF THE GENUS PIERIS IN NORTH AMERICA’
WILLIAM HOVANITZ = ’
1140 ]F. Orange Grove Ave., Arcadia, Calif.
The North American species of the Genus Pieris are distributed
from the Arctic Ocean to Guatemala. However, none of the species
of the genus (sensu stricto) exist in habitats which would be con-
sidered as tundra or arctic-alpine on the one hand, or as wholly tropical
on the other. To this extent, the genus compares somewhat with Colias;
the latter, however, contains two species that do exist in the tundra
habitat.
The concept of a species which has been applied here is the same
as that which has been applied previously to the species of the genus
Colias in drawing the distribution maps for that genus (Hovanitz,
1950). This concept is based upon the consideration that the most
important factor in stability of a species is the genetical population —
or the interbreeding unit. Since, however, genes and their combinations
cannot be seen, nor analyzed except by means of their effects on the
appearance and physiology of the individuals carrying them, they must
be studied through a comparative study of their morphological, anatom-
ical or physiological characteristics. Thus, a name applied to one
population may be applied to another population, or to many popula-
tions, if their characteristics indicate that a similarity exists between
the hereditary makeup of such different populations. Such a name
may cover quite a diversity of differences in the same population, or
even a diversity of different populations if they are genetically separated
one from the other by geographical or ecological means. The extent of
gene interchange as well as the evaluation of the isolation mechanism
determines whether or not one population should be considered as the
’Aided by a grant from tbe National Science Foundation, Washington, D.C.
2The author wishes to thank the following for having made this distributional work possible
by making available the collections under their care: C. B. MacNeil, California Academy of
Sciences, San Francisco; L. M. Martin, Los Angeles County Museum, Los Angeles; T. N. Freeman,
Entomological Research Institute, Ottawa; F. Rindge, American Mus'^um of Natural History,
New York; Harry Clench, Carnegie Museum, Pittsburgh; W. S. Field and J. F. Gates Clark,
U.S. National Museum, Washington, D.C.; and the Dept, of Entomology, Chicago Natural History
Museum, Chicago, Illinois.
3The base maps used in this paper are from Goodes Series of Base Maps by the University of
Chicago Press, used by permission.
73
74
HOVANITZ
/. Res. tepid.
’same species” as another, or a "subspecies” or "geographical race”
of. it, or should not be considered significantly different. These criteria
will be more exhaustively dealt with in a later paper. For practicability,
different criteria must be used at different times and in varying circum-
stances. The ultimate taxonomic designation is a product of weighing
a great number of problems, some factors being known and others
unknown, with the hope that the best result may be achieved. Another
general rule underlies these studies; namely, that stability and uni-
formity are desirable in nomenclature, and that past taxonomic
decisions should not be upset by hasty, poorly-thought-out changes. But
the past should not remain immutable when the decision to change is
demanded by sound study or experimental evidence. Implicit in all
taxonomic studies is perhaps an amount of individual opinion, perhaps
greater than in many more experimental sciences, but even this can be
greatly reduced by conscious effort to separate fact from opinion. It is
the belief of the author that all facts upon which decisions are based
should be stated; these are the "data” which can be refuted. New
decisions should be made then only upon presentations of new data.
Taxonomy by "hunches” is to be avoided; decisions should be
accompanied by evidences for and against any decisions made.
THE SPECIES OF PIERIS
There are five clearly defined species of Pieris in North America.
Of these, only four were native prior to 1800. Besides these, there
are two other groups of related populations which might be considered
"species,” or sibling species, and a large number of geographical varia-
tions within each of these species, many of which are distinctive enough
to be called "geographical races” or subspecies. Division of the species
into geographical races is not to be considered in this paper, with the
exception of the two sibling species just mentioned.
Pieris rapae. This is one of two common European species of
Pieris, the first immigrants* of which appear to have been introduced
into North America prior to I860. Since that time, the species has
spread over the whole of the Unitd States and much of Canada and
Mexico, wherever the natural vegetation has been at least partially
replaced by European weeds, such as black mustard, or by cruciferous
agriculture. This species has a wide range of habitat which is suitable
for its existence, since it is found from the warm parts of the Gulf of
Mexico nearly to the tundra of Canada, and from the Imperial Valley
below sea level to high mountain meadows in the Sierra Nevada or the
Rocky Mountains. The food plants utilized are a variety of cruciferous
plants, among which are all varieties of cruciferous garden vegetables
(cabbage, kohlrabi, kale, etc.) as well as many weedy plants such as
mustards, radish, sisymbrium, etc. Some garden flowers such as
nasturtium are also utilized. In view of the "weedy” nature of this
species, no map is given to illustrate its distribution.
i(i):73-S3, 1962
PIERIS DISTRIBUTION
75
Pieris napi (Fig. 1). This species is found throughout northern
Europe and Asia where there is in effect a continuity with the forms
of North America across the Bering straits and the islands of the
Bering Sea, This is the most northern of all of the species of Pieris,
its range extending to the edge of the tundra or slightly beyond along
its northern limits. The northern distributional limit from the Macken-
zie River and east follows closely the northern limits of the tree line.
The species exists along both the Pacific and Atlantic coast lines, but
extends farther southward on the Pacific Coast than on the Atlantic
Coast. The southern-most population on the immediate Pacific Coast
line is at Lopez Canyon, San Luis Obispo County, California at about
35° north latitude. On the east coast at sea level, the most southern
locality known is along the coast of New York and Connecticut
opposite Long Island at 42° north latitude. However, at increasing
elevations in the Appalachian Mountains southward, P. napi ( or
P. vifginiensis) may be found as far south as the Great Smoky
Mountains of Tennessee and North Carolina at about 35° north
latitude. The southernmost extension of range of Pieris napi in the
strict sense is in the vicinity of Connecticut and the Appalachians of
Vermont, New Hampshire and Massachusetts. Westward, the range
passes through the lower part of Ontario, Michigan, Wiconsin, Minne-
sota, Manitoba, Saskatchewan and Alberta, always staying north of the
prairies. A fuller discussion of the relation of Pieris napi to P.
virginiensis is to be had in a separate paper to follow this one.
Where the range of P. napi reaches the Rocky Mountains in central
Alberta, the species extends clear to the Pacific coast and southward
in the Rocky Mountains at increasing elevations to southern New
Mexico and Arizona. The species appears to be absent in the drier
ranges of the Great Basin and northward in the sagebrush country
into south-central British Columbia. Elsewhere, it is widely distributed
in the habitats that it prefers, these being damp wooded areas with
partial shade, and temperatures not over seventy degrees Fahrenheit..
Of interest in the search for extensions of range are the following
specific locations on the southern parts of the range of Pieris napi:
California: Coast — - San Luis Obispo County, Lopez Canyon,
December through May.
California: Western side Sierra Nevada, Merced County, Yosemite
Valley, 3-4000 feet, April-May.
California: Eastern side Sierra Nevada, Placer County, 6-7000 feet,
June.
Nevada: No records at present.
Arizona: Apache County, White Mountains, August; Gila County,
"Globe” July.
New Mexico: Otero County, James Canyon, Cloudcroft, August,
Catron 6, Mogollon, May, August.
Georgia: No records.
76
HOVANITZ
/. Res, tepid.
FIGURE 1.
Map showing the North
ndpi.
American distribution of Pieris (Pieris)
(i):73-^3, 1962
PIERIS DISTRIBUTION
77
South Carolina: No records.
North Carolina: Black Mountains, April.
Tennessee: Great Smoky National Park, Cherokee, April.
Some other points of interest in the distribution of this species are
the following records :
Pribilof Islands, Alaska, July. Mouth of the Mackenzie River, North
West Territories.
Great Bear , Lake, N.W.T. Churchill, Manitoba ( Hudson Bay ) .
James Bay, Hudson Bay, Quebec. Ramak, Labrador
Pieris napi is the only known butterfly to inhabit the Aleutian
Island^ or the islands of the Bering Sea. This ability is probably
correlated with its adaptation to existence at temperatures below 70 °F,
with low solar radiation and high humidity.
Pieris (Pontia) protodice (Fig. 2). This is the most widely
distributed species of Pieris in North America. Some prefer to place
this species, together with the following two species, in a separate
genus Pontia for which there is much to recommend. To do so,
however, would reduce the size of the. genus Pieris beyond what would
seem reasonable, and, therefore, to obviate this difficulty, and yet show
phylogenetic relationship, Pontia would best be retained as a subgenus
name.
Pieris protodice is best adapted to habitats with more sunshine
than is required by P. napi. It also is enabled to survive in habitats
much too warm for that species. However, cold does not seem to be
a limiting factor as long as there is a short warm season and plenty of
sunlight. These conditions, to some degree, pertain to the area occupied
by the species. It apparently does not flourish in areas of continual hot
temperatures as indicated by its absence in all fully tropical climates.
Pieris protodice differs from P. napi in its absence all along the
immediate Pacific Coast from northern California through Alaska in
keeping with its dislike for cool, cloudy areas. It is absent in the
northern half of Alaska but eastward its distributional limits parallel
those of the timbered area, as with P. napi. East of the Hudson Bay,
however, P. protodice is absent north and east of Ottawa and New
York State. The reasons for this event are baffling unless the present
distribution of the species has been greatly disturbed by man in the
eastern regions of North America as has been found true for Colias
eurytheme. Pieris protodice prefers sunny, open fields as compared
with P. napi which prefers partially shaded, damp areas. The cutting
of the eastern deciduous forest to produce open fields for agriculture
has aided both Colias eurytheme and Pieris protodice in extending their
ranges and population density. It has probably a^so reduced the range
and population density of Pieris napi since the habitats are conflicting.
This probably accounts for the known diminution of Pieris napi in
the region of Cambridge, Massachusetts where they were once exceed-
ingy abundant, and now rare.
78
HOVANITZ
/. Res. Lepid,
FIGURE 2. Map showing the North American distribution of Pieris (Pontia)
protodice.
(0!73-^3,
PIERIS DISTRIBUTION
79
Pieris protodice is found throughout the southeast from Long Island
to the southern part of Florida and around the Gulf of Mexico to
Guatemala. It is very abundant in the highlands of Mexico but may
be found nearly everywhere except in woods or forested areas. It is
found from the southern tip of Lower California northwards, having
a great preference for open areas. It is possible that its great abundance
in southern California coastal areas is a recent event, correlated with
the change of the natural perennial grasslands to annual European
grasses, mixed with Brassica nigra, the black mustard. It reaches its
greatest abundance, however, in the desert regions where conditions
of cool nights, hot days, full sunlight and periodic extensive abundance
of food plants {Cleome, Brassica, Sisymbrium, etc.) allow the species
to swarm in clouds. In such areas, the population abundance is usually
limited by the periodicity of available water.
In the mountain areas, the species is spread from the lowest valleys
to the top of the highest mountains. The author has collected Pieris
protodice at the top of Mt. Whitney, 14,500 feet elevation, the highest
point in the United States south of Alaska, and in the bottom of
Death Valley and Imperial Valley below sea level.
The relationship of Pieris occidentalis to Pieris protodice is here
left in doubt, but for the purposes of the map (Fig. 2), they are
considered forms of one species. This point will be considered in more
detail in a future paper. There is no doubt that at higher elevations
and cooler temperatures, populations exist which have a blacker, fuller
pattern on their wings than populations which exist at lower elevations
at higher temperatures. The preponderance of the evidence suggests
that these differences are not hereditarily controled, but that they are
dependent entirely upon the environmental conditions. Within the
species as a whole, as with P. napi, the darker indiyiduals exist in the
more northern latitudes at cooler temperatures than the lighter ones.
However, this relationship must be reconciled with local factors. In
Colorado, Brown (1957) has shown that P. occidentalis appears to
form definite populations at high elevations whereas P. protodice forms
populations at low elevations. F. Rindge, however, in a personal com-
munications has found both in considerable number in the same valley
in Utah. These points cannot be reconciled at the present time.
P. protodice when exposed to low temperatures develops a color
pattern (more extensive melanin) which is similar to, or identical
with, P, occidentalis. They could very likely be genetically the same
and if they are genetically the same, they are not deserving of a scientific
or Latin name of significance nomenclatorially. This point has not
been proved, however. In the case of geographical variations, such
deviations are considered subspecific unless proved otherwise. In the
case of altitudinal or seasonal differences, such as in Pieris protodice
or Colias eury theme, it is suggestd that Latin names not be applied
except to geographically isolated populations and then in a subspecies
sense only.
80
HOVANITZ
/. Res. Lepid.
FIGURE 3. Map showing the North American distribution of Pieris (Pontia)
sisjmbrii.
(i):73-h> 1962
PIERIS DISTRIBUTION
81
Pieris (Pontia) sisymbrii. (Fig. 3) The Pontia subgenus of the
Pierids all prefer dry habitats with considerable sunshine exposure,
unlike Pieris napi or even Pieris rapae which prefer or can survive in
shade and high humidity. Pieris sisymbrii is one of these. Its habitats
are always open, highly exposed, usually rocky places. It prefers cool
temperatures but with high solar radiation. It lives in the ’spring time
in, the desert, as in the Mohave Desert where it flies at temperatures of
highs not over 70° F, even though the temperatures later in the year
will be over 100° F. It also lives in the mountains at elevations of over
10,000 feet in June and July. In the coastal area of California, its
habitats are usually dry, rocky areas (serpentine, etc.) which are more
dry and more exposed to solar radiation than its general position on
the map would indicate.
Geographically (Fig. 3), the species exists throughout the moun-
tainous areas of the west from the Yukon Territory to Mexico. It
probably also exists in northern Mexico but there are no records.
Outside of this area, there are a series of locations in the Northwest
Territories from the Great Bear Lake southward past the Great Slave
Lake to Lake Athabasca. Between these locations and the Rocky
Mountains there are no locations known. This is probably due to lack
of many suitable habitats. Collectors would do well to increase the
known distribution of this species. It has a wide distribution but is
characterized by the extreme isolated nature of them. Its habitats are
narrowly restricted even on the desert and the adults have a short adult
life, flying for only a few weeks during the year at any one place. The
species aestivates and hibernates as a pupa which is very hard and
impervious to water.
There is relatively little variation between the individuals in a
population in this species, and also relatively little variation geographi-
cally between populations. Some variations have been described, how-
ever, and will be considered in a subsequent paper.
Pieris protodice is a species which is generally distributed and has
many generations per year; it is therefore genetically and physiologi-
cally pliable as is necessary to meet these changed conditions. On the
other hand, Pieris sisymbrii is greatly restricted to isolated habitats, and
the adults fly only a short time; thus, the species genetically may be
said to have developed a narrower range of tolerance through a nar-
rower range of selective factors of the environment.
Pieris (Pontia) heckerii. (Fig. 4). Of all the species of North
American Pieris, P, heckerii has the most restricted geographical range,
extending from south-central British Columbia east of the Cascade
Mountains southwards east of the Sierra Nevada into the deserts of
southern California (Fig. 4). Its eastward limits coincide with the
Rocky Mountain system in Montana, Wyoming and Colorado. The
species does not extend into New Mexico so far as is known, unlike
Pieris sisymbrii. The most southern-known limits are southwestern
82
HOVANITZ
J. Res. tepid.
FIGURE 4. Map showing the North American distribution of Pieris (Pontia)
beckerii.
(i):7}-Sl. 1962
PIERIS DISTRIBUTION
83
Colorado, central Arizona, and southern Nevada, except for California
where the species may be found in the foothills of the mountain ranges
especially on the desert side. However, the species has been found on
the coastal side in the area of the Santa Susana Valley (Moorpark),
San Fernando Valley, Santa Clara River Valley and on the immediate
coast from San Diego and southwards.
Pieris heckerii exists at elevations of sea level at the coast and some
parts of the desert to 9000 feet in the mountain valleys. It has always
been found in areas that are relatively dry or desert-like with high solar
radiation. Its distributional range coincides almost perfectly with that
of the semi-arid brushlands extending from the rain-shadow valleys of
central British Columbia, throughout the Great Basin and inter-moun-
tain valleys of the Rocky Mountains. The factors that limit its distribu-
tion in southern Arizona, New Mexico and northern Mexico are
unknown. Perhaps there is a lack of proper food plant. The distribu-
tion of the species in southern California south of the Great Basin
is correlated closely with the distribution of Isomeris arhorea, its
preferred food plant. It seems likely, however, that some other plant
may also satisfy for this purpose. There is little doubt that intensive
searching for additional poptilations of Pieris heckerii in the range
of Isomeris will extend the distribution some considerable distance,
especially in the south Coast Ranges of California, the foothills of the
Sierra Nevada and southward in Baja California.
Unlike Pieris sisymhrii which has a single short generation per
year, in southern California, Pieris heckerii has a succession of genera-
tions with the bulk of the adult flight being correlated with the time of
of the winter rains. However, in the mountainous regions north of the
Mohave Desert, the species has only one known generation, in the
early summer.
There is little geographical, local population, or seasonal variation
in Pieris heckerii. However, some races have been described and will
be considered in a later paper.
LITERATURE CITED
BROWN, F. M., D. EFF AND B. ROTGER. 1957. Colorado butterflies.
Proc. Denver Mus. Nat. Hist. Nos. 3-7:1-368.
HOVANITZ, WILLIAM. 1950. The Biology of Colias butterflies. 1. The
distribution of the North American species. Wasmann J. Biol. 8:49-75.
J
Journal of Research on the Lepidoptera 1(1) :85-88, 1962
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Copyright 1962
DID THE CATERPILLAR EXTERMINATE
THE GIANT REPTILE?
S. E. FLANDERS
University of California Citrus Research Center and
Agricultural Experiment Station, Riverside, California
The abrupt disappearance of the giant reptile during, the
Cretaceous period brought to an end a very successful plant-animal
system. For 120 million years the reptile population had roamed over
large areas of the earth, subsisting in the lush vegetation covering such
areas. From the standpoint of the reptile s subsistence, plant life was
unlimited. It was an apparently inexhaustible resource.
The possibility that the great reptiles disappeared as a result of
malnutrition and starvation has been considered and discarded as quite
untenable. That highly evolved plant-feeding animals do not normally
become limited in numbers by starvation is axiomatic. Nicholson
(1954) pointed out that the major influence which prevents phytopha-
gous animals from reducing the earth s vegetation to extre'me sparseness
is their attack by natural enemies.
Exhaustibility of plant life, an inherent characteristic of many plant
and animal relations, is rarely realized, being usually precluded through
the regulation of the animal population by its natural enemies ( Flanders
1959). Thus, the Cretaceous populations of the vegetarian reptiles,
by serving as prey for populations of highly efficient predatory reptiles,
were kept at levels well within the limits set by the amount of plant
life. At the same time these predators served also to suppress the
evolution of competitive types of animals of similar size such as the
mammal.
The geological record of the Cretaceous period, however, reveals a
series of events which may have culminated in the temporary destruc-
tion of vast am.ounts of plant life, a period in which the vegetarian
reptiles may have been so devitalized that they either failed to
reproduce or fell easy prey to their predatory relatives.
The critical events which could have led up to the disappearance
of the giant reptile were: (1) the replacement of the ancient fern-
like plants by phanerogamic plants; and (2) the emergence of the
Lepidoptera, an insect order which with few exceptions feeds only
on such plants (Imms, 1930).
The background against which these events should be visualized
85
86
FLANDERS
/. Res. Lepid.
is the world-wide dominance of insects both in species and in popula-
tion densities, a dominance which existed throughout the Age of
Reptiles and the Age of Mammals. The pre-Cretaceous insect fauna
appears to have been quite similar in kind and in numbers to the
post-Cretaceous insect fauna except for the lack of species which fed
on flowering plants (Carpenter 1952). A number of such species were
subsequently derived from the pre-Cretaceous insect orders of Coleop-
tera, Diptera, Hemiptera, Hymenoptera and Orthoptera, species whose
immediate progenitors had presumably already acquired their natural
enemies.
According to Imms (1931), it is practically certain that the Lepi-
doptera is pre-Tertiary in origin, highly organized individuals being
represented in Eocene rocks. The paucity of insects in Cretaceous rocks
is attributed to the absence in that period of suitable fresh-water
deposits in which specimens could have accumulated and become
fossilized.
Plant-feeding insects like other plant-feeding animals acquire
natural enemies which serve to conserve the plant life upon which they
subsist. The fact that the conservation of plant life is a function of
the natural enemies of Lepidoptera has been demonstrated during the
past sixty years by the importation and establishment in North America
of the natural enemies of the caterpillar for otherwise the reduction in
brown-tail moth, Nygmia phaeorrhoea (Donovan), the satin moth,
Stilpnotia salicis (Linnaeus), and the larch caseberer, Coleophora
laricella (Hubner) (Clausen 1956). During the past three years
the writer, by using the grain moth Anagasta and its natural parasite,
Exidechthis, has demonstrated in the laboratory the role of a natural
enemy in the conservation of plant material. In a constant environment
parasitization continuously conserved the plant material despite the
presence of a persistent feeding population of Anagasta.
In nature, therefore, a community of plant-feeding animals con-
sisting, say, of a pentatomid bug, a grasshopper, a moth caterpillar,
a vole, a rabbit, and an ungulate depending on the same resource, grass-
land vegetation, can remain stabilized for a long period of years ( Elton
1946). Such stabilization, however, consists in the total amount of
plant life conserved, not in the relative abundance of the plant species
involved, an abundance determined by the phytophagous animals
through their effect on the competitive capacities of the plant species
(Wilson 1949).
Devastations of plant life by caterpillars can occur today whenever
the regulating action of their natural enemies is disrupted by environ-
mental factors. Graham (1939) reported that during a ten-year period
ending in 1920 an eruption of the spruce budworm, Choristoneura
fumiferana (Clemens), was characterized by flights of moths so
numerous that in the tree tops they had the apparance of a snow storm.
During this eruption the caterpillars destroyed a volume of wood
(i):85-88, 1962
CATERPILLAR AND REPTILE
87
sufficient to supply all of the pulp mills then operating for a period
of forty years.
The capacity of a lepidopteran in the absence of its natural enemies
to reduce great areas of lush vegetation to extreme sparseness and
to maintain it so thereafter was demonstrated in Australia when the
caterpillars of Cactoblastis cactorum (Berg) imported from Argentina
in 1925 destroyed within six years approximately 50 million acres of
the prickly pear, Opuntia spp. Dodd (1929) in reporting on this
devastation stated that great care had been taken not to set free any
of the natural enemies of the catrpillar for otherwise the reduction in
prickly pear would have been of minor proportions.
It is evident that the plant-consuming capacity of a caterpillar
population could equal that of a giant reptile population and that this
capacity because of the caterpillar’s very short life cycle could be
attained at a much greater rate. Supplementing this capacity was a
power of survival much greater than that of the giant reptile, the
minimum food requirements of the individual caterpillar being
infinitesimal relative to that of the individual reptile.
Nevertheless, there is no reason to believe that the caterpillar and
the giant reptile could not have subsisted together on the same
vegetation if the caterpillar, immediately upon its Cretaceous
emergence, had been subject to a regulation by natural enemies as
effective as was that of the vegetarian reptile by its predatory relatives.
The theory that the caterpillar exterminated the giant reptile rests
on the assumption that between the emergence of the new insect order
and its accessions of regulative natural enemies there occurred a brief
period in which the caterpillar population regulated the world’s supply
of plant food, a condition conducive to great variations in abundance
of plant life and of caterpillar life, a condition characterized by "feast
and famine" in which the caterpillar was able to survive but not the
great reptile.
The inherent weakness of the reptile was an extraordinary need
for an abundance of plant material. Only a few years of plant scarcity
could have exterminated it. Hordes of caterpillars, in rapid consump-
tion of fig and breadfruit, laurel and willow, oak and magnolia, could
have so restricted the spatial distribution of the giant reptile that
either its diet was inadequate or it was unable to avoid its natural
enemies.
The small size of today’s descendent reptiles, the vegetarian turtle,
the predatory crocodile, the snake, and the lizard, is evidence of the
giant reptile’s elimination by starvation and predation.
The abrupt end of the Age of Reptiles during the Cretaceous period
is ascribed to a newly emerged order of insects, the Lepidoptera, on
the supposition that for a brief period it regulated the world’s supply
of plant life at starvation levels for the dependent reptiles. Thus, the
giant reptiles which had survived during eons characterized by great
88
FLANDERS
/. Res. tepid.
changes in climate, continental uplifts, and different diets may have
been exterminated by the lowly caterpillar.
LITERATURE CITED
CARPENTER, F. M. 1952. Fossil insects. 1952 U.S.D.A. Year Book.
pp. 14-19.
CLAUSEN, C. P. 1956. Biological control of insect pests in the continental
United States. U.S.D.A. Tech. Bull. No. 113, 151 pp.
DODD, ALAN P. 1929. The progress of biological centrol of prickly pear
in Australia. Commonwealth Prickly-Pear Board, Brisbane Qld. 44 pp.
ELTON, CHARLES. 1946. Competition and the structure of ecological
communities. Jour. Animal Ecol. 15: 54-68.
FLANDERS, S. E. 1959. Biological control. Jour. Econ. Ent. 52: 784-5.
GRAHAM, S. A. 1939. Principles of forest entomology. McGraw-Hill, N.Y.
410 pp.
IMMS, A. D. 1930. A general textbook of entomology. E. P. Dutton & Co.,
Inc., N.Y. 703 pp.
IMMS, A. D. 1931. Recent advances in entomology. P. Blakiston’s Son &
Co., Inc., Philadelphia. 374 pp.
NICHOLSON, A. J. 1954, An outline of the dynamics of animal popula-
tions. Australian Jour. Zool. 2:9-65.
WILSON, F. 1949. The entomological control of weeds. Int. U. Biol. Sci.
Ser.B. 5:53-64.
Journal of Research on the Lepidoptera
1(1) :89-93, 1962
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Copyright 1962
FURTHER EVIDENCE OE THE DISTRIBUTION OF
SOME BOREAL LEPIDOPTERA IN THE
SIERRA NEVADA'
C. H. ERIKSEN
Los Angeles State College, Los Angeles, California
Except where roads penetrate the high country, little or only
scanty information is abaiiable concerning the animal life of the high
Sierra of California. This condition is extant because these mountains
are high, rugged and largely accessible only by trail. The fact also that
collecting gear must be packed in and out encourages few people to
undertake such studies.
Our knowledge of the Lepidoptera fauna of the Sierra stems largely
from the work of Garth (1935) and Tilden (1959). Garth’s study of
the Butterflies of Yosemite National Park is notable since he not only
provides a list of species taken within the confines of the park, but
relates them to the life-zones in which they are normally found to fly.
Tilden has refined the latter concept in his Tioga Pass studies by listing
associations of smaller scope than life-zones.
While backpacking the John Muir TraiP during the summers of
1953, 1954 and 1955, the author made spot collections of Lepidoptera
at various locations along the route (Figure 1). Such was a brief
attempt to add information similar to that of Garth and Tilden to our
knowledge of the day-flying Lepidoptera in high areas from Yosemite
on the north to Mount Whitney on the south. All forms collected and
altimdes at which they were taken are listed in Table 1. To alleviate
any confusion, all nomenclature is after McDunnough (1938).
Altitudes were determined from Starr ( 1953) .
Since Garth (1935) and Tilden (1959) have shown that various
species fly only within certain altitudinal ranges while others are
unrestricted in their flight, there is little need to repeat similar fiindings
here. However, the data collected does extend the known altitudinal
range of several species. The following were taken at elevations higher
than previously recorded and are associated with the life-zone of this
extension.
1 My thanks to Nelson Baker, Santa Barbara Museum of Natural History, for help with identifica-
tion of the material and W. Hovanitz for aid in preparation of this paper.
2The John Muir Trail follows a 215 mile route along the Sierra crest from Yosemite Valley on
the north to Mount Whitney on the south. Except for a short distance out of Yosemite Valley
(elevation 4,000'), altitudes range from approximately 7,000' to 14,I00'.
89
90
ERIKSEN
/. Res. tepid.
Argynnis mormonia (Arctic- Alpine)
Euphydryas chalcedona ( Transition, Canadian )
Lycaena helloides ( Canadian, Hudsonian )
Plebeius aquilo podarce ( Arctic-Alpine)
Plebeius saepiolus ( Arctic-Alpine )
Plebeius icarioides (Hudsonian, Arctic-Alpine)
Plebeius acmon ( Hudsonian )
Garth does not list Lycaena helloides above the Transition zone nor
Plebeius saepiolus above the Hudsonian. Even though Tilden does not
actually say, he intimates that both species transcend all zones. In the
event that any confusion may arise, both species are included in the
above list.
It is of interest that those forms listed in this paper, which also
are found in Colorado, have already been collected in that state from
life-zones here described as extensions (Brown et al, 1957).
LITERATURE CITED
BROWN, F. M., D. Eff and B. Rotger. 1957. Colorado Butterflies. Proc.
Denver Mus. Nat. Hist., Nos. 3-7:1-368.
GARTH, JOHN S. 1935. Butterflies of Yosemite National Park. Bull. So.
Cal. Acad. Sci., 34(1) :l-39.
McDUNNOUGH, J. 1938. Check List of the Lepidoptera of Canada and the
United States of America: Part L Macrolepidoptera. Mem. So. Cal. Acad.
Sci., 1:1-275.
STARR, JR., WALTER A. 1953. Guide to the John Muir Trail and the
High Sierra Region. Sierra Club, San Francisco, 130 pp.
TILDEN, J. W. 1959. The Butterfly Associations of Tioga Pass. Wasmann
J. Biol., 17(2) :249-271.
I (i): 8^-93, 19^2
BOREAL LEPIDOPTERA
91
92
ERIKSEN
/. Re$. Lepid.
Arctic Alpine
Mather Plateau
11,500* ; 7-30-53
ro
€n
Bighorn Plateau
11,500* 1 7-31-54
bH'
ffH
a-
Tyndall Creek
11,000* i 7-29-54
pH
cO
-
Upper Bubbs Cr,
11,000’ ; 7-29-54
sH
Evolution Lake
10,990* s 7-26-53
a>
Hudsonian
Mdl* Palisade Low, L,
10,650* t 7-28-53
sH
Cascade Valley
10,400* 1 7-5-54
Duck Uke Trail
10,100* s 7-5-54
-
tH
Purple Lake
10,000* S 7-6-54
CD
ffH
OI
1000-Island L.
9,850* 1 6-30-54
OI
■H
(tH
Little Pete Mdw,
9,100* s 7-12-55
O
*H
Shadow Creek
9,000* s 7-1-54
(O
(H
CM
Shadow Creek
8,750* 1 7-2-54
Tuolumne Mdws,
8,700* j 6-29-54
CM
pH
Canadian
Palisade Creek
8,125* 1 7-31-55
eH
CO
Quail Meadows
7,700* 1 7-8-54
CO
iH
tH
.H
ra
Table 1.
John Muir Trail Collection
Sites, Listed by Elevation,
and the Species and Number
Taken
Map Location Number 1
Papilio zelicaon Luc,
Papilio rutulus Luc,
Parnassius clodius Men,
>
-o
m
m
Q
X
>.
s,
p
«
m
O
o
Colias behrii Edw,
Pieris sisynbrii Bdv,
Pieris occidentalis Reak,
Argynnis montivaga Behr
>
(0
•H
G
0
1
o
s
m
G
G
<
Brenthis epithore Edw,
c
o
•o
0
0
»H
ns
^ •
o »
m X
r@
$4
^ *
>, >
3 ^
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Euphydryas sierra Wgt,
«
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3
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Melitaea palla Bdv,
Phyciodes laontana Behr
Polygonia zephyrus Edw,
(i); 8^-93. 1962
BOREAL LEPIDOPTERA
93
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CM
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CM
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<0
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pH
CO
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Nymphalis antiopa L.
Junonia coenia Hbn.
Basilarchia lorquini Bdv.
Callophrys dumetorum
Derulexa 3. & B.
Lycaena edit ha Mead
Lycaena helloides Bdv.
lycaena cupreus Edw,
Leptotes marina Reak.
Plebeius aquilo podarce
F. £ F.
Plebeius saepiolus Bdv.
Plebeius icarioides Bdv.
Plebeius acmon West & Hew.
Plebeius shasta comstocki
_E22$ ^ ^
Philotes battoides Behr
Lycaenopsis pseudargiolus
echo Edw.
Thoybes nevada Scud,
Polites sabuleti tecumseh
Grin.
Celerio lineata Fabr.
Pseudohazis hera Harr.
Anarta cordigera Thun.
1(1) :95-96, 1962
Journal of Research on the Lepidoptera
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Copyright 1962
ARGYNNIS AND SPEYERIA
BY WILLIAM HOVANITZ
Department of Zoology ^ Los Angeles State College
The use of names for nomenciatorial purposes involves judgment
on two levels. The first level involves nomenciatorial laws, and the
second is the judgement on the part of the taxonomist as to the most
reasonable and useful application of a name for a biological category.
When a name is being considered for use on the generic level,
the first point is its nomenciatorial availability. Here, it is important
that the name not be a homonym, and that it have priority according
to the established laws of nomenclature. A species is designated as the
type species of that genus. For example, the species paphia L. has been
designated as the type species of the genus Argynnis.
Once the type species has been established for a genus, and there
are no doubts of the nomenciatorial status of either, the specific name
or the generic name, no one who wishes to remain within established
criteria for nomenciatorial uniformity and fairness has any right to
change this name. The generic name is strictly applied, however, only
to the one species. It is the prerogative of any individual to use the
same generic name for other species if he deems that the species
concerned ought to be considered congeneric. Strictly speaking, it is
possible for each species in existence to be the type species for a
different generic name. If each of these names were used, each species
would be in a different genus and the beneficial effects of binomial
nomenclature would be rendered worthless.
The purpose of a binomial nomenclature is to indicate relationships
by grouping related species into the same genus. How this should be
done is the prerogative of the individual taxonomist. It is assumed
that such a person will use the method that is best suited to his purposes
in showing relationships.
The genus Argynnis in the broad sense is composed of many
species, some of which can be grouped into categories having biological
similarities, but which are smaller in content than the genus
when considered with a broad view. Generic names have been used
for some of these smaller categories with various species designated as
types. It is perfectly satisfactory from a nomenciatorial as well as a
biological standpoint to do this. This is the situation with regard to
the use of Speyeria. The type species of the name as a genus is idalia.
95
96
HOVANITZ
/. Res. tepid.
For those persons who wish to consider idalia as not congeneric with
paphia, or with any other older name, Speyeria is a valid generic name.
Or, it can be used in a subgeneric sense, in which case Argynnis would
again be used as the generic name.
This author believes that the broad use of the genus Argynnis is
preferable to the practice of restricting the name to the paphia group
of species, and further believes that the genus can be divided into some
more or less satisfactory subgenera of which Speyeria is one. Our
American species then would be designated Argynnis {Speyeria) idalia
to show its relationship. The other American Argynnids then would
also continue to use the name Argynnis rather than Speyeria.
It is not the purpose of the editor of this journal to insist on any
particular terminology for to do so would impinge on the freedom
of the scientific worker, who alone has the right to make his choice.
This editor can only insist on sound data to back the decisions of
authors in their use of terminology, as in drawing conclusions from
their work.
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Volume 1
Number 1
August^ 1962
IN THIS ISSUE
Editorial 1
Variation in the silvering of Argynnis (Speyeria) callippe in
the interior mountain area of south central California . . .
Oscar Elton Sette 3
The effect of various food plants on survival and growth rate of
Pieris .... William Hovanitz and Vincent C. S. Chang 21
General characteristics of the movements of Vanessa cardui (L.)
J. W. Tilden 43
Three factors affecting larval choice of food plant
William Hovanitz and Vincent C. S. Chang 51
The generic, specific and lower category names of the Nearctic
butterflies. Part 1 — The genus Pieris . . Paddy McHenry 63
The distribution of the species of the genus Pieris in North
America William Hovanitz 73
Did the caterpillar exterminate the giant reptile? ....
S. E. Flanders 85
Further evidence of the distribution of some boreal Lepidoptera
in the Sierra Nevada Clyde Eriksen 89
Argynnis and Speyeria William Hovanitz 94
FRONT COVER ILLUSTRATION
PARALLEL COLOR VARIATION IN SOME CALIFORNIA NYMPHALIDS
Column 1, E.chalcedona
Column 2, E.editha
Column 3, C.leanira
Column 4, C. pal la
Column 5, C.hoffmanm
Details in Ecology 22: 259-284, 1941.
Ci>
E
@F RilSEARCHJ
THE LEFI
Volume 1
Number 2
]a^ary, 1963
Established in 1 962
Published at
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Edited by WILLIAM HOVANITZ
WITH EMPHASIS ON ENVIRONMENTALLY AND GENE-
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ogy, geographical variation, speciation, physiology, etc. In short,
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Journal of Research on the Lepidoptera 1(2) :97-108, 1963
1140 ’IF. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 196}
COMPOSITION AND RELATIVE ABUNDANCE
IN A TEMPERATE ZONE BUTTERFLY FALMA
THOMAS C. EMMEL and JOHN F. EMMEL
Reed College, Portland, Oregon md Stanford University, California
The Conner Pass area, lying on the crest of the Sierra Nevada
range in Placer County, California, provides a rich variety of habitats
lying between 6900 and 8300 feet elevation. During the summers of
1956 and i960, the authors made an intensive study of this region and
we have reported elsewhere (Emmel & Emmel, 1962a, 1962b) on
the ecology and factors affecting distribution of the 74 species com-
posing the butterfly fauna. We wish to report here the interesting data
obtained on the faunal composition and relative abundance of species
within the seven major Rhopalocera groups recorded in the Donner
Pass area, and to evaluate the possible factors influencing this distribu-
tion of species.
INTRODUCTION
Hovanitz (1958) has. recorded the occurrence of Rhopalocera
families and genera at each five degrees of latitude and 1000 meters
of elevation above sea level for the entire New World. However, this
study did not include the numbers of species or genera in each family
that occurred in each of these areas. Tilden (1959) divided the butter-
flies of the Tioga Pass area under five plant associations; Emmel &
Emmel (table 1, 1962a) partitioned the Donner Pass fauna under
four plant associations. But these authors did not compare the faunas
of each association or attempt to ascribe reasons for the greater number
of species of a particular family in one association than another. The
purpose of this paper is to report an examination of the composition
and habitat associations of the Donner Pass fauna and from this
examination to derive some understanding of the factors affecting the
distribution and success of species in a butterfly family within a
given set of environments.
DESCRIPTION OF STUDY AREA & METHODS
In a previous paper (Emmel & Emmel, 1962a) ^ four general
vegetational associations representing the union of floras from three
97
98
EMMEL AND EMMEL
/. Res. Lepid.
life zones (Transition, Canadian, and Hudsonian) were delineated,
and may be briefly reviewed.
1. The W et-M.eadow habitats (elevation 7000-7260 feet) support
verdant expanses of grasses (Poa species) and sedges (Car ex), Lilium,
Mimulus., Delphinium, Castilleia, and other wildflowers, with scattered
thickets of willows (Salix) along the stream banks. Trifolium, Viola,
Aster, Potentilla, and other lepidopterous food plants (see table IV)
are common.
2. Dry-Meadoiu areas (elevation 6800-7600 feet) are moist in
the spring but are dry throughout July and August. Fireweed (Epilo-
bium angustifolium) , Achillia, Calyptridium, and Penstemon, as well
as grasses, are typical plants. Food plants include Kibes, Polygonum,
Lupinus, Gnaphalium, and others as listed in table IV.
3. Dense Forest (elevation 6800-7600 feet), mainly of Canadian-
Zone trees, covers about one-half of the total Pass area. Red Fir
(Abies magnifca), Pinus contorta, P. monticola, and Mountain Hem-
lock (Tsuga mertensiana) represent the majority of the forest trees.
Only three known butterfly food-plant genera are found here: Pinus,
Arceuthobium, and Ceanothus,
4. The Montane areas (elevation 7160-8383 feet) are generally
open talus slopes, where the dry, almost soilless environment discourages
the growth of most trees. Included in the flora of these mountain peaks
are Rabbit Brush (Chrysothamnus) , Sage Brush (Artemisia), Mule-
Ears (Wyethia mollis), Dipinus, and Sedum species. Eriogonum,
Astragalus, Potentilla, Ceanothus, and other food plants in Table IV,
are widely distributed here.
Thus the diversity of vegetational associations in this temperate-
zone locality provides habitats and food plants for a considerable
number of butterfly species, and it is of interest to compare the relative
success of species in different families in colonizing this favorable yet
rather small area (six square miles) in the Sierra. In 1956 and I960,
the number of species (see Emmel & Emmel, 1962a, for a detailed
listing) in each of the four vegetational zones was determined by
intensive collecting throughout Donner Pass.
In the course of studying the influence of meteorological conditions
on the flight activity of these species in I960, daily counts of the
numbers of flying butterflies were made within the 500 x 800 feet
"Lodge Meadow” study area (see Emmel & Emmel, 1962b), which
contained 46 species recorded for the Pass (see Table III). This area
is composed of both wet and dry meadows. These counts were made
by direct observation and represented numbers seen for each species
between 10 and 12 a.m. daily. The total number of butterflies re-
corded in this latter study was 7,720. To determine "success” in
numbers of flying adults, the daily counts for numbers of individuals
for each of the species in a family were added together, and this figure
was calculated as a percentage of the total for all families. Thus the
contribution of each family toward the composition of the Donner
(2):97-!oS, 196}
DONNER PASS BUTTERFLIES
99
Pass butterfly fauna can be compared on a percentage basis, both as
regards numbers of species and numbers of individuals.
OBSERVATIONS
Some 74 butterfly species were found in the Donner Pass area,
and these species belonged to seven of the major families of Rhopalocera
found in the western United States. The number of species in each of
these groups varied from 1. fof the Danaidae to 28 for the Lycaenidae,
or in terms of percentage of the total Pass fauna, 1.4% to 37.8% of
ail the species. Table I shows the composition of the Rhopalocera groups
recorded in the total Pass area.
Table II and Figure 1 show the composition of the butterfly fauna
recorded in each of the four general habitats of the Donner Pass area;
the figures in Table II represent the per cent of total species in each
habitat that belong to each family, while the graphs in Figure 1 show
actual number of species for each family in each habitat. It is imme-
diately obvious that the montane and dry-meadow habitats have the
largest faunas of the four areas, and that the Lycaenidae family is usually
dominant as regards number of species.
As stated earlier, it was possible to obtain some data on the number
of individuals of each species (46 in all) occurring in the Lodge
Meadow study area. The composition and relative abundance of the
six major Rhopalocera groups recorded there are given in Table III.
For the most part, there is a good correlation between number of species
and number of individuals for each family.
DISCUSSION AND CONCLUSIONS
The number of butterfly species found in this six-square-mile area
of the Sierra Nevada is unusually high. Tilden (1959) records only 43
species as occurring in the Tioga Pass region, representing a study area
of approximately the same size although of somewhat higher elevations
(over 9941 feet above sea level). Garth (1935) records about 100
species for all of Yosemite National Park (area 1,179 square miles,
elevations from 2000 to 13,090 feet), and the present authors and
Lloyd M. Martin (personal communication) can note from their
experience that the average restricted Sierran locality seems to support
around 35 to 65 species. Thus the Donner Pass area is particularly
interesting from the standpoint of invesigating the reasons for the
occurrence and abundance of a species in a specific habitat, for as noted
earlier (Emmei & Emmel, 1962a) this region is a meeting place for
a variety of habitats- and a high number of plant species, and these
factors may be viewed as possible reasons for this variety of Rhopa-
locera.
The relative proportions in number of species among the Rhopa-
locera groups involved seems to follow the relative number as found
throughout North America north of Mexico (determined from Ehrlich,
1961 ). The descending order of these groups in terms of number of
100
EMMEL AND EMMEL
/. Jl«, Lepid.
TABLE I. Composition of the seven major Rhopalocera groups
recorded in the Donner Pass area.
Family
No, of Species
% of Total Species
Papilionidae
5
6,8
Pieridae
8
10,8
Danaidae
1
1,4
Satyridae
2
2.7
Nymphalidae
20
27,0
Lycaenidae
28
37,8
Hesperiidae
10
13,5
TOTALS
74-
100,0%
"In our earlier paper (Emmel 5 Emmel, 1962a) ,
74 species and 2 "forms”, Anthocaris sara reakirti and
Colias eurytheme amphidusa, were listed, making a total
of 76 phenotypically-distinct entities
DONNER PASS BUTTERFLIES
101
species is (both in the continent and local Pass faunas): Lycaenidae,
Nymphalidae, Hesperiidae, Pieridae, Papilionidae (follows Satyridae in
the continent fauna), Satyridae, Danaidae.
However, when the total Pass fauna is divided in terms of occurrence
in each of four habitats, patterns of apparent preference for certain
habitats emerge — both for all families combined and for individual
groups (Table II and Figure 1). The highest total number of species
(60) occur in the montane habitats, the next highest (55) in the dry
meadow areas, the third highest (44) in the wet meadows, and the
lowest number (26) in the forest areas. The descending sequence of
environmental support of number of butterfly species is thus:
Montane > Dry Meadows > Wet Meadows > Forest.
This pattern, obtained by comparing numbers of Rhopalocera species,
is probably explained by the observed fact that the montane areas
contained a greater variety of ' mico-habitats” than any other area. These
micro-habitats included small stream areas, exposed mountain tops,
open and brushly talus slopes, etc., ranging from 7100 to 8383 feet in
elevation, the greatest range in altitudes for any of the four "macro-
habitat” areas considered. Such variation in ecological conditions
(especially from edaphic and climatic standpoints) permits a wide
variety of potential food plants to flourish. Axelrod (I960) has
commented on the marked diversity of habitats in montane areas
and the possible influence of this condition on early angiosperm evo-
lution through permitting rapid development and adaptive radiation.
Such diversity also influences butterfly distribution, variation, and
speciation (see Emmel & Emmel, 1962a; Le Gare & Hovanitz, 1951;
Moeck, 1957), and provides potential areas for successful invasion by
butterflies already occurring elsewhere.
The large number of total species in the dry meadow areas is
believed to be due to the great variety of known food plants occurring
there; the wet meadows supported correspondingly fewer food-plant
species. The jorest areas contained the least variety of micro-habitats
and food plants; also, solar radiation, an important influence on butter-
fly flight habits (see Emmel & Emmel, 1962b), was obviously at a
considerably lower level in this habitat than in the other three. Table
IV shows the number of known food-plant genera in each habitat for
each Rhopalocera group. These and the preceding data agree with the
theory that a greater variety in each of these Sierran butterfly groups
is more likely to occur where micro-habitats and food plants are most
varied, as these conditions, together with a high level of solar radiation,
provide support for a wider selection of species than in such areas as
pine forests.
This theory can be applied equally to the question of why one
group has more species than another group in a certain habitat. The
Papilionidae are more successful in forest than in montane areas in
terms of per cent of total species there, but they actually have the
most species in the latter areas. So we must consider:
102
EMMEL AND EMMEL
/. Res. Lepid.
WET MEADOWS DRY MEADOWS FOREST
MOr^TANE
4
PAPiLION/DAE
6
4
2
PIERIBAE
DANAIDAE
Is;:;
SATYRIDAE
HESPERUDAE
Fig. 1. Numerical composition of the butterfly fauna recorded io each of
four general habitats of the Donner Pass area.
the
NUMBER OF SPECIES
i(2):^7-io8, 15163
DONNER PASS BUTTERFLIES
103
1) That habitat in which the group has its highest percentage
of the total species population:
Papilionidae Forest {1.1%)
Pieridae Forest (15.4%)
Danaidae Forest (3.8%)
Satyridae Montane (3.3%)
Nymphalidae Forest (38.4%)
Lycaenidae Wet Meadows (45.5%)
Hesperiidae Forest (19.2%)
2) That habitat in which
number) :
Papilionidae
Pieridae
Danaidae
Satyridae
Nymphalidae
Lycaenidae
Hesperiidae
the group has the most species
Montane (4)
Dry Meadows, Montane ( 6 )
( 1 in each habitat)
Montane (2)
Montane (16)
Dry Meadows ( 22 )
Montane (10)
( direct
The really significant consideration in evaluating a group’s contribu-
tion to the fauna in a series of habitats would seem to be the number
of species it has existing in each habitat. Of course, a further and
perhaps more important consideration would be the number of indi-
viduals of the group in that habitat, but these data were not obtained.
However, speaking from the standpoint of number of species the
butterfly groups are uniformly most successful in the montane and
dry meadow habitats.
As opposed to evaluating the most suitable habitat for most groups,
one can also consider the success of individual groups in colonizing
or existing in the various habitats. In the wet meadow, dry meadow,
and montane habitats, the Lycaenidae are the most abundant group — •
with 20 to 22 species in such areas. The Nymphalidae take second
place (10 to 16 species) except in the forest zone where they lead the
other groups in number of species. The Hesperiidae are third — with
5 to 10 species in each zone — except for being second in the forest
areas (leading the Lycaenidae). The Pieridae are uniformly fourth
(4 to 6 species) while the Papilionidae have only 2 to 4 species in
each zone. There is only one species in the Danaidae (all habitats,
however) and two in the Satyridae (montane habitat only).
This apparent relative success of each group, in our opinion, is not
due to a "better” adaptation (the result of natural selection) of a
group’s species in one or another habitat, as Ehrlich (1962) has
emphasized. Instead, from our observations, success is likely due to
the fact that food plants of species in the seven Rhopalocera families
were more varied (as to number of kinds; Table IV), generally more
numerous (as to number of individuals of each plant species), and
104
EMMEL AND EMMEL
/. Res. Lepid.
TABLE II. Composition of the butterfly fauna recorded in each of the
j.
four general habitats of the Donner Pass area.
Family
Habitat
Area““
Wet Meadows
Dry Meadows
Forest
Montane
Papilionidae
4.5
5.5
7.7
6.7
Pieridae
9.1
10.9
15.4
10,0
Danaidae
2.3
1.8
3.8
1.7
Satyridae
0.0
0.0
0.0
3,3
Nymphalidae
27.3
27.3
38.4
26.7
Lycaenidae
45.5
41.0
15.4
35.0
Hesperiidae
11,4
14,5
19.2
16.7
(1) TOTAL SPECIES
44
55
26
60
(2) PERCENTAGE OF
TOTAL PASS FAUNA
(74 species)
REPRESENTED:
59.5%
74.3%
35.2%
81.1%
“The numbers in each vertical column (file) represent percent of
total species in that one habitat (the total number of species is found
at the bottom of each column). The actual numbers of species of that
family in each habitat, from which the preceding percentages were
calculated, are shown in graphic form in Figure 1,
““See Emmel S Emmel (1962a) for map and complete description of
all habitats in the Pass area.
i(2);97-to8, 196}
DONNER PASS BUTTERFLIES
105
more generally distributed for the Lycaenids than for the Nymphalids,
for Nymphalids than for Hesperids, etc. It is concluded in this paper
that the composition of a butterfly fauna is affected by the require-
ments (or adaptation) of each butterfly species in that fauna for a
certain food plant, in addition to a certain level of solar radiation and
other factors as noted in Emmel & Emmel, 1962a. In turn, food plants
obviously have grater chances of finding proper environmental condi-
tions in a macro-habitat that contains a number of varied micro-habitats,
each with particular edaphic and climatic conditions. The greater the
number of host food plants, the greater the number of potentially
successful butterfly species, and this theory is believed to account for
the observed distribution of the 74 butterfly species (of seven major
groups) among the four macro-habitats of the Donner Pass area.
TABLE III, Composition and relative abundance of the six major Rhopalocera
groups recorded in the Lodge Meadow habitat.
Family
No, of Species
% of Total Species
% of Total
Individuals
Recorded
Papilionidae
2
4.4
4.7
Piaridae
4
8,7
9,8
Dana i da e
1
2.2
0.3
Nymphalidae
15
32,6
44.9
Lycaenidae
16
34.8
23,2
Hesperiidae
8
17.4
16,7
TOTALS
46
100.1%
99.6%
SUMMARY
1. The Donner Pass area in Placer County, California, supports
an extraordinary number (74) of Sierran Rhopalocera species, and
its varied habitats and vegetational associations make the area par-
ticularly interesting for investigating the basis for the occurrence and
abundance of a species and a family in a specific habitat.
2. The number of species in the seven major families of the Pass
area are proportionately equivalent to the numbers of species in these
106
EMMEL AND EMMEL
/. Res, Lepid.
TABLE IV, Number of known food plant genera in each habitat
for each
Rhopalocera family"
Rhopalocera
Habitat
Area
Family
Wet Meadows
Dry Meadows
Forest
Montane
PAPILIONIDAE
Salix
—
Ceanothus
Cymopterus
Salix (stream)
Ceanothus
Prunus
Sedum
FIERI DAE
Cruciferae
genera
Cruciferae
genera
Pinus
Cruciferae
genera
Trifolium
Trifolium
DANAIDAE
-
-
-
SATYRIDAE
( Grasses )
(Grasses)
(Grasses)
Grasses
NYMPH ALI DAE
Viola
Castilleia
Salix
r’lantago
Castilleia
Aster
SlrsEum
Ribes
Gnaphalium
Lupinus
Ceanothus
Castilleia
Aster
Chrysopsis
Ceanothus
Salix (stream)
Lupinus
LYCAENIDAE
Salix
tupinus
Ceanothus
Lupinus
Arceuthobium
Pinus
Ceanothus
Lupinus
trifolium
Potentilla
Polygonum
Ribes
Eriogonum
Potentilla
Calyptridium
Polygonum
Sedum
Eriogonum
Astragalus
HESPERIIDAE
Potentilla
Grasses
Sidalcea?
Potentilla
Grasses
Sidalcea
—
Potentilla
Grasses
"Data summarized from text of Eramel 6 Emmel, 1962a.
(2):97-Jo8, 1963
DONNER PASS BUTTERFLIES
107
groups as found for the whole of North America (north of Mexico).
From most to least number of species: Lycaenidae, Nymphalidae,
Hesperiidae, Pieridae, Papilionidae (follows Satyridae for North
American fauna ) , Satyridae, Danaidae.
3. The highest total number of butterfly species in Donner Pass
occurs in the montane habitats ( 60 species ) . The dry meadow habitats
support 55 species, the ivet meadow habitats have 44 species, and the
jorest areas have 26 species occurring in them.
4. The basis of this distribution is explained by the theory that
a greater food-plant diversity occurs in the areas (such as the montane
macro-habitats) that have a number of micro-habitats, each with
particular edaphic and micro-climatic conditions, and that this host-
plant diversity promotes the immigration and continued successful
existence of more Rhopalocera species than in less diversified macro-
habitats ( such as these Sierran forest habitats ) . The possible influence
of such factors as solar radiation and climate acting directly on butter-
flies rather than plants is also considered.
5. Data obtained on the number of flying individuals of all species
found in the Lodge Meadow area showed there was usually a good
correlation between number of species and number of individuals for
each family in this restricted Sierran locality.
SYSTEMATIC LIST OF RHOPALOCERA OCCURRING
AT DONNER PASS, PLACER COUNTY, CALIFORNIA
PAPILIONIDAE
1. Papilio zelicaon Lucas
2. Popilio indra indr a Reakirt
3. Papilio rutulus Lucas
4. Papilio eurymedon Lucas
5. Parnassius clodius baldur
Edwards
PIERIDAE
6. Neophasia menapia
Felder & Felder
7. Pieris sisymbrii Boisduval
8. Pieris protodice Linnaeus
9. Pieris rapae Linnaeus
10. Euchloe creusa hy antis Edwards
11. Anthocaris sara form julia
Edwards
12. Colias eurytheme Boisduval
13. Colias phUodice eriphyle
Edwards
DANAIDAE
14. Danaus plexippus Linnaeus
SATYRIDAE
15. Ceononympha tullia calif ornica
Westwood
16. Cercyonis sthenele oetus
Boisduval
NYMPHALIDAE
17. Speyeria cyhele leto Behr
18. Speyeria zerene zerene
Boisduval
19. Speyeria coronis snyderi Skinner
20. Speyeria atlantis irene Boisduval
21. Speyeria mormonia arge Strecker
22. Boloria epithore Edwards
23. Chlosyne palla Boisduval
24. Chlosyne hoffmanni hoffmanni
Behr
25. Phyciodes campestris montana
Behr
26. Phyciodes mylitta Edwards
27. Polygonia zephyrus Edwards
28. Nymphalis calif ornica Boisduval
29. Nymphalis milberti Latreille
30. Nymphalis antiop a Linnaeus
31. Vanessa cardui Linnaeus
32. Vanessa atalanta Linnaeus
33. Vanessa virginiensis Drury
34. Vanessa car ye Hubner
35. Precis lavinia Cramer
36. Limenitis lorquini Boisduval
& Leconte
LYCAENIDAE
37. Satyrium calif ornica Edwards
38. Satyrium sylvinus Boisduval
39. Satyrium saepium Boisduval
40. Satyrium behrii Edwards
41. Satyrium fuliginosa Edwards
42. Strymon melinus Hubner
43. Callophrys johnsoni Skinner
44. Callophrys nelsoni Boisduval
108
EMMEL AND EMMEL
/. Res. Lepid.
45. CaUophrys augustinus iroides
Boisduval
46. CaUophrys eryphon Boisduval
47. CaUophrys dumetorum perplexa
Barnes & Benjamin
48. Lycaena arota virginiensis
Edwards
49. Lycaena edit ha Mead
50. Lycaena nivalis Boisduval
5 1 . Lycaena cupreus Edwards
52. Lycaena heteronea Boisduval
53. Everes comyntas amyntula
Boisduval
54. Plebejus anna Edwards
55. Plebejus saepiolus Boisduval
56. Plebejus icarioides Boisduval
57. Plebejus shasta Edwards
58. Plebejus acmon Westwood
& Hewitson
59. Plebejus {acmon?) lupini
Boisduval
60. Agriades glandon podarce
Felder & Felder
61. Glaucopsyche lygdamus hehrii
Edwards
62. Philotes enoptes Boisduval
63. Philotes battoides intermedia
Barnes & McDunnough
64. Celastrina argiolus echo
Edwards
HESPERIIDAE
65. Thorybes nevada Scudder
66. Pyrgus ruralis Boisduval
67. Pyrgus communis Grt.
68. Erynnis juvenalis Fabricus
69. Erynnis afranius Lint.
70. Hesperia juba Scudder
71. Hesperia nevada Scudder
72. Hesperia harpalus Edwards
73. Polites sonora Scudder
74. Polites sabuleti tecumseh Grin.
ACKNOWLEDGMENTS
The authors again express their appreciation to Mr. William N. Goodall
of the National Audubon Society, Director of the former Audubon Camp of
California at Donner Pass (Norden, California), for extending full coopera-
tion to us during the collection of the data reported in this series of three
papers on the Donner Pass butterfly fauna. Drs. Thomas Harvey and Kenneth
Tanksley of the Camp staff kindly furnished plant identifications. Discussions
with Mr. Lloyd M. Martin, Associate Curator of Entomology at the Los
Angeles County Museum, have illuminated a number of points included in
these papers. The authors wish to thank Dr. G. Frank Gwilliam, Associate
Professor of Biology at Reed College, for his help in preparing the present
paper.
LITERATURE CITED
AXELROD, DANIEL 1. I960. The evolution of flowering plants. In The
Evolution of Life, Sol Tax, ed. Pp. 227-305. Univ. of Chicago Press.
EHRLICH, PAUL R., & A. H. 1961. How to Know the Butterflies. Brown,
Dubuque, Iowa. 262 pp.
— — — - — ■, & RICHARD W. HELM. 1962. Patterns and populations.
Science 137: 652-657.
EMMEL, THOMAS C., & JOHN F. EMMELL. 1962a. Ecological studies of
Rhopalocera in a High Sierran community — Donner Pass, California.
I. Butterfly associations and distributional factors. Journ. Lepid. Soc.
16: 23-44.
— — , 1962b. Ecological studies of Rhopalocera in a High
Sierran community — Donner Pass, California. II. Meteorologic in-
fluence on flight activity. Journ. Lepid. Soc., in press.
GARTH, JOHN S. 1935. Butterflies of Yosemite National Park. Bull.
So. Cal. Acad. Sci. 34: 37-75.
HOVANITZ, WILLIAM. 1958. Distribution of butterflies in the New
World. In Zoogeography. American Association for the Advancement
of Science, Publication No. 51, Washington, D. C. Pp. 321-368.
LE GARE, MARY JUDE, & WILLIAM HOVANITZ. 1951. Genetic and
ecologic analyses of wild populations in Lepidoptera. 11. Color pattern
variation in Melitaea chalcedona. Wasmann J. Biol. 9" 257-310.
MOECK, ARTHUR H. 1957. Geographic variability in Speyeria. Mil-
waukee Entomological Society. 48 pp.
TILDEN, J. W. 1959. The butterfly associations of Tioga Pass. Wasmann
J. Biol. 17: 249-271.
Journal of Research on the Lepidoptera 1(2) : 109-1 13, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 1963
THE ARGYNNIS POPULATIONS OF THE SAND
CREEK AREA, KLAMATH CO., OREGON
Part I: The effect of the formation of Mt, Mazama on the area
AND ITS POSSSIBLE INFLUENCE ON THE BUTTERFLY FAUNAS OF THE
Sand Creek Basin.
J. W, TILDEN
San Jose State College, San Jose, California
INTRODUCTION
The populations of Argynnis (Speyeria) found in eastern
Oregon share with similar populations in many parts of western
United States, a tendency to great variation and to a breakdown of
the distinctive appearance of many of the named geographical pheno-
types (subspecies). The region drained by Sand Creek, lying east of
Crater Lake in Klamath County, Oregon, is occupied by populations
of Argynnis exhibiting this variation to a marked degree.
The Sand Creek area was chosen for study not only because of
the variability of its Argynnis fauna, but in particular because it is
a region the recent geological history which is fairly well understood,
and inferences may be drawn of the possible effect of this geological
history on the several species of Argynnis now inhabiting this region.
Part I of this paper will deal with the recent geological history of
Mt. Mazama and the nearby lands to the east of Crater Lake, and the
possible influence of these events on the faunas now living in this
region. Part II will attempt an analysis of the variation of these
Argynnis populations, and is postponed until much more field work
can be done so that a proper appreciation of the extent of variation
is possible.
BACKGROUND
The stability of certain isolated populations has been interpreted
as due to long occupation of the present habitat. This time element
has been construed as allowing stabilization, without undue influence
from surrounding populations. Such conditions infer stability in the
habitat, without the disturbing influence of earth changes in the area.
Such stable habitats have been referred to as refugia.
Opposed to such a condition of stability, many portions of the
earth s surface have been subjected to changes more frequently, and
some of these changes have occurred in very recent times. Eastern
Oregon is a region of many lava flows, and scoria and cinder falls, and
109
110
TILDEN
/. Res. Lepid.
the effects of these actions are still clearly visible in numerous places.
It is suggested that such phenomena must have had a major effect
upon any butterfly populations existing in the regions where such
events took place.
THE SAND CREEK BASIN AND ITS
RECENT GEOLOGICAL HISTORY
Sand Creek rises at Anderson Spring, elevation 7088 ft, slightly
northeast of Kerr Notch in Crater Lake National Park, and runs
easterly, leaving the park in the near vicinity of the now unused East
Entrance. It continues out of the park, the elevation gradually decreas-
ing. The area chosen for study is that from Lost Creek Campground,
Crater Lake National Park, 5972 ft., located between Sand Creek and
its branch, Wheeler Creek, and the point at which Highway 232
("Old 97”) crosses Sand Creek. This area is readily accessible. Work
in the Crater Lake National Park was possible under a permit from the
National Park Service.
The Argynnis samples indicate that the following species are repre-
sented: Argynnis {Speyeria) coronis Behr, zerene Bdv., callippe Bdv.,
egleis Behr, atlantis Edw., and hydaspe Bdv. Possibly others may be
present in lesser numbers but have not yet been collected. It is inter-
esting to note that Argynnis cybele {leto) appears to be absent from
the region, and also from all of Crater Lake National Park, though
common west of the Cascade divide, and also in the Blue Mountains
of eastern Oregon. The scarcity in the Sand Creek Basin of suitable wet
habitats, or the recency of earth surface changes, may be clues to its
absence.
Fair samples of these various species, collected by the author and
also by others, have been examined, and it is evident that the degree
of variation is surprising. As stated above, analysis of these and future
samples is postponed until later, but enough information is now avail-
able to make it apparent that the populations of each species involved
are far from stable. Many individuals resemble no named geographic
segregate. Some species are represented by individuals resembling two
or more named segregates and it is evident that no equilibrium in
phenotype has been reached by any of the several species inhabiting
the area.
Crater Lake is not a true crater, but a caldera, brought into its
present form by the collapse inwardly of the summit of Mt. Mazama.
In its most developed state, Mt. Mazama was more than 12,000 ft. in
height. During the Pleistocene, it developed its greatest elevation,
produced a great lava flow, and was subsequently glaciated extensively.
The effects of this glaciation are visible around the present rim of the
lake. Later, the present Crater Lake was formed by the collapse inwardly
i(2):io^-Jij, 196}
SAND CREEK ARGYNNIS
111
of the upper mile or so of Mt. Mazama, with attendant glowing
avalanches of scoria and pumice and great clouds of pumice dust.
The area east of the lake was covered with pumice and scoria from
the glowing avalanches. The land still further east was covered by
pumice falls that reached a depth of ten feet or more close to the
mountain, and gradually thinned out easterly, extending in at least a
very thin layer, approximately fifty miles to the east.
It is believed that the glaciers, or some of them, must still have
been present when the collapse took place, since the residue of scoria
and pumice is light in certain areas around the rim. This would be the
situation if the scoria and pumice fell onto glaciers and was carried
away in part. If it had fallen in every case on bare ground, it would
be left there. This may explain why the present pumice soils are more
extensive in the non-glacial areas. It is believed also that the collapse
took place in winter, because there is relatively little evidence of
severe forest fires in the wake of the collapse. If the collapse had taken
place in the summer, it has been reasoned that the forest fires surely
would have been more extensive.
There seems to be agreement that most or all of the vegetation was
destroyed within the effective range of the scoria and pumice avalanches
and of the heavier pumice fall. If this is correct^ the present vegetation
of the entire region must have regrown since the collapse took place.
The rate of recovery of the vegetation seems to have been inversely
proportional to the depth of the pumice and scoria that covered a
given area. The prevailing winds are believed to have been much the
samx then as now, basically western, since the pumice was carried much
further east than west.
The Sand Creek Basin is situated so that it received very heavy
pumice fall, and very probably, extensive to complete vegetational
destruction.
Carbon dating of wood that is imbedded in the pumice, and which
is therefore assumed to have been charred by the glowing avalanches
of pumice and scoria, sets the time of the collapse at 6640 ± 250 years.
In round numbers, popular accounts state this as 6550 years. From
this it is inferred that the Sand Creek Basin became a sterile habitat
at that time.
It is of interest to observe that such streams as Upper Sand Creek
and Wheeler Creek, below Kerr Notch, have deep V-shaped canyons,
geologically very young and without sign of glaciation such as that
found on the rocks of the Rim above. From this it seems clear that
Sand Creek acquired its present bed recently. Indeed, its present course
may very well be consequent to the recent geological events of the
area. The depth of the pumice is also clearly visible.
Knowledge of the date when the vegetation was destroyed allows
some speculation concerning present vegetational cover of the region.
Recovery of the vegetation must have lagged for many years. The
types of vegetation found in the total Crater Lake region permit some
112
TILDEN
/. Res. Lepid.
understanding of the relative times of recovery. In Crater Lake National
Park, the forest along the southern boundary is a mature Sierran Coni-
ferous Forest of Ponderosa Pine, Sugar Pine, White Fir, Douglas Fir
and Incense Cedar, with an understory of Snow Brush {Ceanothus
velutinus) and Greenleaf Manzanita {Arctostaphylos patula) . The
forest of the flanks of the mountain on the southern and western slopes
is a mature Mountain Hemlock-Shasta Fir Forest, with increasing
amounts of Subalpine Fir at higher elevations. Such forests have
relatively less understory. The highest elevations support Whitebark
Pine.
Northerly and easterly, in the areas of greater pumice fail, (as for
instance the Pumice desert) the forest has not even yet succeeded in
taking over the denuded and pumice-covered land. In the intermediate
areas where the forest is slowly invading and colonizing the .pumice
soils, the successful pioneer is Lodgepole Pine, which on such soils
forms a singularly uniform and barren type of forest with very few
underplants.
The soils of the study area are formed of pumice and scoria and
from the foregoing evidence, are derived from the pumice fall from
the collapse of Mt. Mazama. The forest here is essentially Lodgepole
Pine Forest, of the type found also on the pumice at higher elevations
in the park. The soils and the forest are evidently contemporary in the
Sand Creek Basin, with those occurring on pumice within the park
itself. We should regard the forest cover of the Sand Creek Basin as
equally recent.
Dating of the pumice fall and the destruction of the vegetation
give a reference point. Vegetational recovery must involve a very
considerable period of time. This reasonably will be longer in areas
where the pumice fall is deeper. While no exact figure can at present
be suggested for the age of the current forest cover of the Sand Creek
Basin, it must be very much less than 6640 ± 250 years. Very likely
portions of it may be a fraction of this age.
From this it seems evident that the Argynnis populations of the
Sand Creek Basin (and similar areas) are occupying habitats that
are the opposite of refugia. The time in occupation of contemporary
populations on such disturbed areas is brief.
DISCUSSION
Assuming an area from which vegetation has been destroyed, with
attendant soil changes, and with sufficient time elapsed to allow new
vegetation to occupy the area, we may well expect certain things to
take place, among which the folowing may be suggested.
1. The vegetation that recolonizes the land will in all possibility
be different than that which previously grew in the area. Most likely
many previous floral components would now find the area inhospitable.
2. Recolonization of the habitat by organisms would theoretically
(2):i09-ii3, 196}
SAND CREEK ARGYNNIS
113
be possible from any portion of the surrounding regions. It may be
expressed by saying that recoionization could be expected to be muUi-
dkectional unless some factor prevented recolonization from one or
more directions.
3. Such recently recolonized areas would contain mixed populations
the components of which may have moved in from any of the sur-
rounding regions.
4. If such heterogeneous populations are studied before adequate
time for stabilization has passed, it is predicted that great variation in
individual phenotype will be present.
Expressing this in terms more directly related to the Argynnis
populations of the Sand Creek Basin, it is tentatively concluded that
the variability of these several populations is in part a result of the
recency with which they have established themselves in this area follow-
ing a major change in habitat, and in part a result of recolonization by
an unknown number of varying populations from the surrounding
regions.
Less tangible are two other possibilities. One is that the time during
which the Sand Creek Basin was unsuitable for habitation by Argynnis
was of sufficient duration to allow the surrounding populations which
it separated, to diverge to some, extent. The other is that the recent
forest cover of the pumice soils may differ sufficiently in its selective
factors io that the eventual stable population may when developed
differ from those around it.
Thanks are due to the National Park Service, and in particular to
Park Naturalist Bruce Black, for extensive courtesies and helpful
information.
REFERENCES
BRIGGS, LYMAN J. 1962., When Mt. Mazama lost its top. National
Geographic, 122 (1): 128-133.
MOECK, ARTHUR H. 1957. Geographic variability in Speyeria. The
Milwaukee Entomological Society, 48 pp.
WILLIAMS, HOWELL. 1961. Crater Lake — The story of its origin.
University of California Press, Berkeley & Los Angeles. XII + 98 pp.,
10 pits., 8 figs.
Journal of Research on the Lepidoptera 1(2) 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 7563
CATERPILLAR VERSUS DINOSAUR?
THEODORE H. EATON, JR.
Museum of Natural History and Department
of Zoology, University of Kansas
A PAPER BY S. E. Flanders in the first issue of The Journal of
Research on the Lepidoptera (Flanders, 1962) bears the intriguing
title, "Did the caterpillar exterminate the giant reptile?” In his dis-
cussion, the "abrupt end of the Age of Reptiles during the Cretaceous
period is ascribed to a newly emerged order of insects, the Lepidoptera,
on the supposition that for a brief period it regulated the world’s
supply of plant life at starvation levels for the dependent reptiles.”
The argument for this hypothesis rests in part on modern examples
of large-scale devastation of certain species of plants by the larvae of
Lepidoptera, and in part on assumptions concerning the time of origin
and expansion of this order, the nature of plant life during the Cretaceous
period, and the habits and relationships of the "giant reptile.”
Many explanations have been proposed for the extinction of dino-
saurs and several other groups of reptiles at the end of the Cretaceous,
but there is not yet any general agreement as to the most important
causes. The proposals are conjectures, having little or no direct evidence
to support them. The theory under discussion adds another to the list.
The consensus among paleontologists now is that one or two "causes”
are not sufficient, and that the extinction of these animals is a more
complicated phenomenon than it has seemed. While it is possible that
destruction of food-plants by caterpillars has at times contributed to
extinction of some herbivorous reptiles, and thus indirectly to that of
their predators, the present writer thinks that a close look at what
happened in the Mesozoic might be helpful in judging this hypothesis.
Our knowledge of fossil Lepidoptera is' scanty, but the earliest
known moth appears to be Loses triassica, described from Upper Triassic
beds in Queensland (Tindale, 1945). It is known only from a fore and
hind wing, of which the venation agrees broadly with that of Jugatae
(Homoneura); Tindale placed it in a new, more primitive suborder,
Eoneura. Thus the order must have originated more than a hundred
million years before the extinction of dinosaurs, although leaf-eating
caterpillars might not have characterized the earliest genera and families
of moths. Reports of moths and butterflies from the Jurassic litho-
graphic limestone in Bavaria are based on specimens now attributed
114
i(2):ii4-ii6, ip6j
CATERPILLAR VS. DINOSAUR?
115
to other orders, especially Homoptera. Unless more information appears
in Frank M. Carpenter s forthcoming volume on fossil insects in the
"Treatise on Invertebrate Paleontology," we can say nothing specific
about Lepidoptera of the Jurassic and Cretaceous; none are mentioned
in the "Traite de Paleontologie,” vol. Ill (1953). In the Cenozoic,
however, moths and butterflies of modern families are known, apparently
from the Eocene on.
It is reasonable to suppose that the rise of Lepidoptera coincided
with that of flowering plants. These plants are known first from the
Jurassic, but at least sixteen modern families are represented in Lower
Cretaceous rocks; by the late Cretaceous many existing species of trees
had appeared. The length of the Cretaceous was about 70 million years.
From its close to the present time was a little less, say 63 million years,
according to recent determinations (Kulp, 1961). But we cannot, in
the nature of the case, show evidence that caterpillars appeared in great
numbers in the late Cretaceous, or that they were then so free of para-
sites or predators that they could, by worldwide devastation of all sorts
of plants, bring starvation upon the reptiles.
Some of the statements of Flanders concerning reptiles bear comment
from the viewpoint of a paleontologist. For instance, "The inherent
weakness of the reptile was an extraordinary need for an abundance of
plant material . . . The small size of today’s descendent reptiles, the
vegetarian turtle, the predatory crocodile, the snake, and the lizard, is
evidence 'of the giant reptile’s elimination by starvation and predation.’’
A sauropod, hadrosaur, ankylosaur or ceratopsian dinosaur may have
eaten a large quantity of vegetation, the amount depending on size and
activity. These animals were present in the late Cretaceous and occupied
a position comparable in some ways to that of the browsing mammals
of the Cenozoic. But their extinction does not seem to have been abrupt;
it was a slow, uneven decline probably extending through millions of
years, and does not differ so far as we know from the decline and
disappearance of various other groups of animals at earlier and later
times. The flying pterosaurs, whose food was probably fish, declined
and died out at about the same time as the dinosaurs. So also did
mosasaurs, a specialized line of giant marine fish-eating lizards; so too
the very different aquatic plesiosaurs, as well as a few late Cretaceous
ichthyosaurs and carnivorous dinosaurs.
No living reptiles are directly descended from any of the above-
mentioned Mesozoic reptiles. Turtles and crocodiles originated in the
Triassic and have continued with no fundamental changes to the present
time. Lizards are known first from the Jurassic, snakes from the Creta-
ceous, and their size cannot be taken as evidence of any changes among
the giant reptiles. These giant reptiles were clearly not a single natural
group, nor, on the other hand, were all members of the orders to which
they belonged of great size. In the two orders of dinosaurs, for example,
there were some species, both early and late, that failed to reach the
size of an average alligator.
116
EATON
/. Res. Leptd.
Efforts to explain the extinction of dinosaurs and various other
reptiles in the Cretaceous period ought to take into account a number
of possible causes. The one proposed by Flanders does not give the
answer to several problems, such as the disappearance of various un-
related and ecologically different groups, not necessarily of large
animals, and the survival beyond the Cretaceous of certain other
reptiles, not necessarily small. There is no evidence of either a biological
or a geological catastrophe of large proportions at the close of the
Mesozoic, unless the slow retreat of epicontinental seas in some areas
can be so described. Probably the answer will eventually be found in a
combination of many factors.
LITERATURE CITED
FLANDERS, S. E. 1962. Did the caterpillar exterminate the giant reptile?
J. Res. Lepid., 1 (1): 85-88.
KULP, J. LAURENCE. 1961. Geologic time scale. Science, 133 (3459) :
1105-1114.
TINDALE, N. B. 1945* Triassic inseas of Queensland. Rroc. Roy. Soc.
Queensland, 56 (5): 37-46.
Journal of Research on the Lepidoptera 1(2) :117-123, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 196}
GEOGRAPHICAL DISTRIBUTION AND VARIATION
OF THE GENUS ARGYNNIS
I INTRODUCTION
IL ARGYNNIS IDALIA
WILLIAM HOVANITZ
California Arboretum Foundation, Inc., Arcadia, California, and
Los Angeles, State College, Los Angeles, California
L INTRODUCTION
The Genus Argynnis in the comprehensive sense covers a wide
variety of form and size in Nymphalid butterflies. The geographical
distribution of the genus as a whole is somewhat comparable to that
of Colias but is actually wider in some areas due to a tropical group
in the eastern hemisphere. A natural desire to have a greater similarity
of type within a "genus” and a desire of some workers to "split” to the
ultimate has left the genus in a diverse state of fragmentation. This has
been aided by regional preferences and the difficulty for most Lepid-
opterists to observe and study members of the group from continents
other than their own. The members of the genus from the smaller
isolated continents such as North America and South America have
apparently either diverged slightly from the type on the largest land
mass (Eurasia) or they have retained more primitive features. In
North America only two of the several subdivisions of type within the
Argynnis appear ever to have arrived for colonization. These two have
diverged to such an extent on that continent that residents of that
continent of natural inclination should consider them as of different
genera {Boloria and Speyeria) . One of these {Speyeria) is only slightly
different from one of the species groups of Eurasian Argynnis, namely,
aglaja, adippe, niohe, etc. and placed by some in a genus Brenthis which
is not the usage of Brenthis used in North America by others.
The variety of the genus in Eurasia is much greater than in North
America. It is natural to expect that the types present in Eurasia which
would be most like those now present in North America would be
those whose climatic adaptations would be the most similar to the
conditions present in the area where a land bridge exists now or did
exist in the immediate past. This is the area of Alaska and adjacent
Siberia. The Boloria group is the best adapted to the climates of this
area and may even now be considered for all practical purposes to have
"continuity” across the barrier at the Bering straits. With the possi-
bility of gene flow not very remote, it is not difficult to see why Boloria
117
118
HOVANITZ
/. Res. Lepid.
of North America and Asia (Europe) should be much alike. In fact,
several species of Boloria can and do go under the same species name
on both continents, just as do three species of Colias which have the
same geographical relationships to the land bridge.
The species which are the most remote in their climatic preferences
from the climates of the Bering area are those species which are the most
divergent on the two continental areas; these are the most southern
types. Argynnis paphia is the member of a southern subgenus of
Argynnis in Eurasia and this subgenus apparently never colonized
America. Instead, a type similar to aglaja apparently did colonize
America and from this presumably all the American Argynnids of
the larger {Speyeria) type subsequently were derived. It is an hypo-
thesis that the most extreme types in North America, such as idalia,
diana and nokomts, which bear considerable pattern and color relation-
ship to Eurasian species such as paphia, sagana, childreni, etc. were
derived independently from them from aglaja — like ancestors under the
selective influence of the comparable conditions of climate to which
they were subjected (hot-humid summers). Comparative correlations
of the facies of the butterflies with climates at different parts of the
world will be made later in this series.
The Argynnis of South America bear no close relationship apparent-
ly to the Argynnis of any other part of the world. There is no easy way
at the present rime for them to have arrived there by migration from
other continents. Only two alternate proposals seem possible. The first
of these is that these Argynnis were isolated many years ago (the early
Tertiary) from the other Argynnis of the world and that they arrived
there by migration from North America at a time when North America
was inhabited by a more primitive type. They might also have arrived
from Africa early in the evolution of the genus should there be reason
to believe seriously in the past movement of South America away from
a more close proximity to Africa. The second proposal on the origin
of the South American Argynnis is one of independent origin in South
America from some tropical or subtropical relative of the North
American Argynnis. This proposal seems very weak in view of the lack
of any close relative that would seem to be a likely ancestor. Dryas
(Colaenis) , Dione and Euptoieta seem to be out of the question, though
this may be only based upon superficial appearances. They feed upon
similar plants, a factor greatly favoring the argument, especially with
regard to Euptoieta which inhabits cold country in the Andes, feeds on
violets and has much the same habits as Argynnis itself.
The question comes up as to what is a genus. Should the entire
group which has been pictured here be considered one genus, or should
it be raised up to a sub-family or even a family. If the latter, then
should the major groups be raised from subgeneric level to the generic.^
If so, then how many groups at this level should be recognized? The
question is then condensed to one problem. There really is no biolo-
gical entity which can be known as a genus. All hopes to the contrary,
(2);ii7-I23, 1963
ARGYNNIS DISTRIBUTION
119
the genus is solely a matter of convenience in nomenclature, which
should show as much as possible, phylogenetic relationships. The
system should start at the bottom of the classification heirarchy. If it is
known what should be considered species, sufficient related species
ought to be put into a genus to make a reasonable group on the basis
of numbers and morphological (genetical) similarity. Should too few
species be put into a genus the advantages of the binomial nomen-
clature are destroyed, in much the same way as would occur if every
person had the family name Smith. A further criterion, in addition to
size and uniformity, is geographical coverage. It is a natural phenome-
non that isolation leads to differentiation. Related members of a species
group may diverge slightly due to continental isolation and yet in all
other ways be closely knit as a single group. The decision here must
rest on the needs of the taxonomist in showing relationships and the
usefulness to biologists. Nomenclature bears one major value: that of
usefulness. If different generic names were to be used on each con-
tinent, a degree of provincialism would develop which would be hard
to penetrate; local butterfly books in North America and in Europe
have now diverged to such an extent in terminology that the only way
a novice can determine relationships between the butterflies of Europe
and North America is to look at the specimens — -an ironic turn of
events since that should be the purpose of nomenclature. As a result
therefore, it is the hope that there may be a return to the larger genera
of the past; this is the thesis upon which the use of Argynnis here is
based, without necessarily accepting the assumption that the generic
limits and the characters used for their delimitation should remain as
in the past if more specific future work should show the need for
change.
IE ARGYNNIS (SPEYERIA) ID ALIA
This representative of the North American Argynnis differs in
appearance from the usual Speyeria probably more than any other
except possibly for A. diana. The details of its pattern which are dis-
tinctive and different from the standard Argynnis pattern are the greatly
increased black pigment around the borders of the wings, even extending
to the basal section and nearly covering the hind wings, but at the same
time leaving free of black pigmentation, the marginal and submarginal
rows of spots ( fig. 1 and 2 ) . The general facies therefore is distinctive and
different from any other Argynnis. At the same time, the black pattern
elements of the center of the fore wing are reduced rather than ex-
panded as in all other related Argynnis. It therefore shows a develop-
ment of a pattern not only unlike that of the Argynnis, including the
subgenus Speyeria as indicated by Dos Passos and Grey, but a trend
in a diametrically opposed direction. The distinctiveness of this species
is such as to need no real description or detailed analysis. Likewise, from
the study of specimens throughout the geographic range of the species,
there are no geographical variations apparent, though there are in-
120
HOVANITZ
/. Re$. Lepid.
Fig. 1. Argynnis idalia, upper side; top: male; bottom: female, from Oak
Park, Illinois, July 8, 1906. F. S. Daggett.
i(2):ii7-i2h 19^3
ARGYNNIS DISTRIBUTION
121
ilfjflllliilililigF
iisSllfiL
iSisiSssSssilism
immmmmmmmmmmm
|bs::ss;s::l
^^asms:
g^^SSSSSSSi
rfw slow*******
W ^Ms^sssi
Fig. 2. Argynnis idalia, same as fig. 1 only lower side.
122
HOVANITZ
/. Res, Lepid,
SCALE
1000 FOOT CONTOUR
Fig. 3. Map of North America showing the distribution of Argynnis idalia.
(2):ii7-i2}, 1963
ARGYNNIS DISTRIBUTION
123
dividual differences in size and slight differences in the pattern observed
on an individual basis. Perhaps statistical differences could, and will,
be detected in the future.
The distribution of this species is limited to a central and eastern
location across the United States and a small part of Canada. The
biological reasons for this restricted distributional range are not known.
It can be seen from the map (fig. 3) that the species ranges from New
England (specimens which I have seen come from as far as Massa-
chusetts though records indicate Portland, Maine as the furthest point
reached north and east), southward to North Carolina (probably erro-
neous records indicate Georgia), and west to Colorado and North
Dakota (probably erroneous records indicate Montana). There are
records or indications in old literature for Arkansas but I have seen
no specimens from there. The species seem to prefer the area south of
the Canadian life zone, that is, the Transition life zone of sorts, though
it is pretty hard to outline any definite climatic zonation. Records
indicate that it is to be found in most of the southern counties of
Michigan and the southern counties of Minnesota. Scudder (1889)
writes: "This butterfly belongs to the Alleghanian fauna, though its
distribution appears to be somewhat irregular. It inhabits lowlands and
is much more abundant in the extreme eastern portion of its range than
elsewhere, unless it be the western prairies.” Mention of it in Louisiana
by Strecker is most probably in error.
A further discussion of the range of this species will be deferred
until the maps of other species have been published, which will be
used for comparison.
REFERENCES
DOS PASSOS, C. F. AND L. P. GREY. 1947. Systematic catalogue of
Speyeria (Lepidoptera, Nymphalidae) with designations of types and
fixations of type localities. Amer. Mus. Novitiates No. 1370.
— — — — 1945. A genitalic survey of Argynninae (Lepidoptera, Nymph-
alidae). Amer. Mus. Novitates No. 1296.
MACY, R. W. AND H. H. SHEPARD. 1941. Butterflies (of Minnesota).
Univ. Minn., Minneapolis.
MOORE, SHERMAN. I960. A revised annotated list of the butterflies of
Michigan. Occasional Papers of the Museum of Zoology, University of
Michigan. No. 617.
SCUDDER, S. 1889. The Butterflies of the Eastern United States. Cambridge,
Mass.
Journal of Research on the Lepidoptera
1(2) :124-134, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 196}
THE RELATION OF PIERIS VIRGINIENSIS EDW.
TO PIERIS NAPI L.
SPECIES FORMATION IN PIERIS?
WILLIAM HOVANITZ
California Arboretum Foundation, Inc., Arcadia, California, and
Los Angeles State College, Los Angeles, California
In a previous paper (Hovanitz, 1962), it was indicated that
a fuller discussion of the relationship of Pieris napi L, to Pieris virgin-
iensis Edw. would be forthcoming. In that paper, the map showing the
distribution of P. napi included the distribution of P. virginiensis but
a closer distinction between the two was not indicated because of the
reduced size of the map included. A more detailed map showing the
relationships between the two biological entities under discussion is
included with this paper, together with a description of the known
facts of the lines of separation of the two "species” or races, and a
summary of the events that have most probably occurred in relation
to the two groups up to the present time.
On the map in the previous paper (Hovanitz 1962), the range
of Pieris napi L. for the eastern United States included virginiensis as
well as napi itself. The race of napi with which we are concerned here
is known as oleracea. This race blends northwards and westwards into
other races by imperceptible gradations. Pieris napi oleracea (known
also by the summer brood names, aestiva Edw., hyemalis Edw. and
acadica Edw.) may be considered the name for the race of napi east
of 90° Longtitude and south of 50° N. latitude, more or less, with the
exception of the area south of New York state (38° N). Other names
can be found in McHenry (1962). The area south of New York
state and east of 90° Longitude is occupied by the race or species
virginiensis. In New York state and other points in the vicinity, or close
to the southern limits of oleracea, there are populations which in some
cases appear to be more closely related to virginiensis than to oleracea,
and in other cases, are more closely related to oleracea with occasional
individuals similar to virginiensis. The status of the line of contact
between oleracea and virginiensis is not one of free and even blending
such as might be expected of freely interbreeding individuals in a case
of solely geographical isolation. Populations are rather clearly either
oleracea or virginiensis as the case may be, with only occasional examples
of intermediates giving some hint of the nature of introgression at the
line of contact.
124
(2) -.124-1 34. 19^3
PIERIS NAPI RACES
125
The contact of these two species or races bears a close relationship
to the similar situation between the species or races Basilarchia astyanax
and B. arthemis in the same general area (Hovanitz 1949).
Specimens of P. napi studied in various collections have permitted
a map to be drawn showing the detailed distribution of oleracea and
virginiensis (fig. 1). On this map, locations where oleracea (or napi in
the species sense) are known are shown by the letter N and virginiensis
locations are shown by the letter V. In addition to the indications on
the map, there are reasons for assuming that each of these races overlaps
the other even greater than seems indicated on the map. Due to lack of
specific data (e.g. "New York”), some of these indications could not
be shown on the map. These details will be mentioned below.
On the map, it can be seen that oleracea extends southwards in the
mountains of New England to Connecticut where it comes in contact
with virginiensis or something like virginiensis . It is doubtful that these
specimens are truly virginiensis, though they do bear some resemblance
to them. Instead they are more likely abberant oleracea, which are
genetically tending in the direction of virginiensis through introgression.
In addition, the habitat change out of the Canadian forest typical of
oleracea to the "Transition” lowlands might lead to a phenotypic
alteration in facies. Populations of considerable size are definitely known
at the present time in Vermont and New Hampshire. They are not now
known in the vicinity of Boston, Mass, though they were common there
before 1850. The following bears witness to a reduction in distribution
of napi in the 1800 s:
From Scudder (1889) :
"This butterfly is a member of the Canadian fauna . .
"It is found throughout New England although seldom abundant south
of the annual isotherm of 48°. Northward and eastward it is everywhere abund-
ant and it continues to be so as far south as Williamstown, Mass. (Scudder),
Dublin, N. H. (Leonard) and Portland, Me. (Scudder, Verrill). South of 42°
N. L. or the northern boundary of Connecticut, it rarely occurs, although it
has been taken at Farmington, Conn. (Norton), Newport, R. I. (Miss
Coggeshall teste Higginson) and Fire Island Beach, Long Island (Smith).
Even in the north, however, there seems to be some local causes for its
abundance.”
"Mr. Lintner, writing in 1864, says that in Schoharie, N.Y., 'it was
extremely rare until recently,’ and previous to 1857, 'he had taken it but
once in three years . . "Mr. Bacon of Natick, Mass, says that the insect by
no means disturbs cabbages and turnips as it did fifteen or twenty years ago.”
"I recollect once seeing the college yard in Cambridge, I think it was
about 1857, fairly swarming with P. oleracea. It is now never found, to my
knowledge, anywhere in the region about Boston, and I think it is wholly
confined to the less cultivated and especially the hilly distrias of New England.”
FromKlots (1951):
"Napi has . . . suffered a great restriaion in habitat, and probably in
habits, for it is now almost entirely limited to shaded Canadian Zone forest
. . "Not recorded south of the Catskill Mountains in New York”.
126
HOVANITZ
/. Res. Lepid.
From Klots (1935) of Edwards,
"He mentions a female taken by Mead in the last week of June at Stony
Clove in the Catskill Mountains . . . many eggs were obtained . . . from the
chrysalis an oleracea emerged.” "Stony Clove is in excellent territory for napi
{oleracea) but I know of no records of virginiensis from there.” "I have . . .
seen specimens of virginiensis from Big Indian Valley in the Catskills, where
the environment is more suited to virginiensis.”
"In 1931, near McLean, Tompkins County, New York, I was able to
obtain undoubtedly authentic eggs, larvae and chrysalis of virginiensis, through
watching females ovipositing on Dentaria diphylla Michx.”
From Fiske ( 1901 ) :
"Pieris napi. once common enough to be injurious to cabbages and other
cruciferous plants throughout New England, has since the introduction of rapae
become one of our rarest butterflies. Its range is now restricted to the northern
and mountainous portions, and even in the fortresses of the White Mountains it
is a scarce insea.”
The eighteenth century idea of a "species” contributed a lot to a
general confusion of the nature of the relationship between oleracea and
virginiensis. It was then, and too often now, the general idea that a
species was represented by a specific morphological type, without regard
to the variation which a population may have within itself. Some of
the best collectors and breeders of Lepidoptera have therefore become
confused, having the all pervading desire to designate some biological
unit as either a "species”, subspecies or variety when the true relation-
ship may not be possible with only these tools of nomenclature. For
example, Edwards (1881) says:
"Virginiensis . . . has become a true species, although unquestionably, in
a higher latitude, it appears as an occasional aberration only of Oleracea.”
Also, Klots (1951) says:
"Until very recently, virginiensis has been confused with napi, but it is
now known to be a distinct species of more southern (Transition Zone)
distribution.”
Edwards (1868-1872) says:
"In the Kanawha district, it replaces Oleracea which is yet unknown there.
It is not uncommon in the month of May, frequenting open woods rather than
gardens, and in this respect differing in habit from the allied species. I have
never met with it later than June, though Oleracea, in the Northern States is
most abundant after that month and continues breeding till the early autumn
frosts.” "I have received specimens of Virginiensis from Mr. Wm. Saunders of
London, Canada and am informed by him that it is there a rare insect.”
DIFFERENCES BETWEEN P. NAPI OLERACEA AND
P. NAPI VIRGINIENSIS
The primary visible differences between P. napi oleracea and P.
napi virginiensis can be observed in figs. 2 and 3. In figure 2 is shown
the male and female of the upper side of virginiensis to the left; at the
same location in figure 3 is shown the under side. These specimens are
(2):i24-n4, 196}
PIERIS NAPI RACES
127
Fig. 1. Map showing the distribution in the eastern part of North America
of the introgression area of Pieris napi races virginiensis and oleracea.
from near Pittsburgh, which although toward the north end of the
range of virginiensis are quite typical of the populations south through
the Appalachian mountains. The pattern of the upper side female
should be noted as quite typical of this race, and which is only rarely
present in oleracea. Of critical distinction, the pattern of the under side
of the wings should be noted. The veins are outlined in a tan pigmen-
tation, not heavy but rather broad in expanse, in appearance unlike
any other race of napi.
P. napi oleracea is shown in figures 2 and 3 by the central two
specimens; these are from southern New Hampshire which is near the
128
HOVANITZ
J. Res. Lepid.
« 2
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6 w
^ On
..U
o §
a M
.. B
Ph S
H •
m
0\
X
§
^ a
^1 S
,3 3
V N
S m
^ G\
G ^
P rn
a ph‘
O G\
H
1 ^
dl ^
e§
^ e«
^ «
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(2);i24-134> 19^3
PIERIS NAPI RACES
129
Same as fig. 1, but all lower side fades'
130
HOVANITZ
/. Res. Lepid.
southern part of the distributional range of this species. They are how-
ever typical in appearance. Note the difference between the two races
on both the upper and lower surfaces: virginiensis is more of chalky
white on the upper side as compared with oleracea; the wings on the
upper side tend to be veined with black near the borders and cell in
oleracea which is replaced with a brownish ''rapae-ty^t' pattern in
virginiensis; on the under side the veins are heavily lined with black in
oleracea in a way distinctive from that of virginiensis; and there is a
yellowish suffusion on the under side of the wings in oleracea not
present in virginiensis. On the other hand, the appearance of oleracea
changes during the seasons while that of virginiensis does not. There is
no indication that virginiensis has more than the one generation per
year, this occurring in early May whereas oleracea has a succession of
generations starting in April and extending throughout the year until
the cold weather of the autumn. During this extended period the weather
conditions are quite different, and the effects of this on the appearance
of oleracea are profound. The appearance of the adults emerged during
the warmer part of the season are quite different. A specimen of the
summer brood is shown in figures 2 and 3 at the upper right. Note the
slightly increased size but more important the almost complete absence
of any pigment other than white on the wing surfaces. These summer
forms have been given the names hyemalis, acadica and aestiva by
Edwards for the general area under consideration. As a racial name, the
only one of any significance however is oleracea. Throughout the ranges
of these races, there is fairly good uniformity in pattern and character-
istics; however, it should be pointed out that within each of these races
and especially oleracea there is a great amount of variation which tends
to blend the two together. For example, a specimen at the bottom right
in figures 2 and 3 is from a population of oleracea but shows the "rapae-
type” pattern of virginiensis. The underside of the wings however makes
clear that the specimen is oleracea.
Areas of typical oleracea may be described as follows: The Can-
adian forest areas ranging from northern Michigan across northern
Ontario and Quebec into eastern Quebec, New Brunswick, Nova Scotia,
Maine and the mountainous parts of the New England states as far as
Massachusetts. South of these points and until the typical populations of
virginiensis are reached in New York state, Ohio and Pennsylvania^ the
populations apparently are in a state of uncertainty regarding a complete
divergence from the virginiensis or the oleracea genome and all the
things pertaining thereto, including in addition to pattern differences,
the physiological alterations required for the more southern existence,
the univoltine life cycle and the changed food plant preferences. These
intermediate populations lie in a line ranging from Michigan, across
southern Ontario to Montreal in Quebec and southward to the Catskill
Mountains of New York. Some of these can be distinguished as oleracea
with virginiensis tendencies and others as virginiensis with oleracea
tendencies, though the former outrank the latter. Populations which
(2);i24-134, 1963
PIERIS NAPI RACES
131
would be considered oleracea with virginiensis tendencies would be the
following, with an indication of the extent of the tendency:
Michigan: Emmett Co., north part of southern peninsula (slight)
Ontario: region around London, Huntsville, Sydney Field station,
Marmora, Trenton ( considerable )
— - — : Aylmer, Bells Corners, Brittania, South March, Merivale,
Constance Bay, Ottawa West (slight)
Quebec: Harrington Lake, Gatineau Park (slight)
Massachusetts: Cummington, Whately, Franklin Co., Mt. Greylock
Populations which would be considered virginiensis with oleracea
tendencies would be:
Quebec: He Perrot (Montreal)
New York: the Adirondacks, the Catskills, Ilion, McLean
Ontario: Hamilton
Populations with little sign of intergradation on the northern end
of the range of virginiensis'.
Pennsylvania: Scranton, New Brighton, Pittsburgh, Forbes Forest,
Charlevoi, Showville, Washington Co., Fayette Co., Lawrence Co.
Ohio: Canton
West Virginia: Forks of the Kanawha, Coalburgh
HABITS OF THE RACES
The 'rather distinct break between the geographical ranges of
oleracea and virginiensis would seem to need some isolation barrier for
the maintenance of its integrity. This barrier is clearly not one of water
or mountain but rather one of habitat selection and the accompanying
biological changes in the physiology of the races concerned. For some
idea of the field habits of these races, we may refer to other authors:
Klots ( 1951 ) says of virginiensis'.
"Until very recently, virginiensis has been confused with napi, but it is
now known to be a distinct species of more southern (Transition Zone)
distribution. Its dark markings are a more smoky, diffuse brown and it lacks
the yellowish tint beneath, on the HW and apex of the FW, which napi
usually has. It is single brooded. More even than napi, it is limited to woods
and very local.”
"Food: — Toothwort {Dent aria diphylla) , probably other Dentaria. One
brood. Adults in early May (central New York.”
"Range:— Transition Zone, Ontario, central New England and New York,
s. to Virginia (TL. Kanawha, West Virginia)
He says of napi oleracea'.
'^Napi has . . . suffered a great restriction in habitat, and probably in
habits, for it is now almost entirely limited to shaded Canadian Zone forest
... A virginiensis lacks yellowish beneath on the HW and apex of the FW
and has the scaling along the veins of the HW and costa and apex of FW
beneath much more diffuse and powdery looking. A rare dark form of P. rapae
lacks dark spots above, but never has the dark-lined veins of napi, and has
heavier scaled, less fragile looking wings.”
Food. Formerly, at least, nearly any cultivated or wild cruciferae; now
probably chiefly limited to native, woodland species, such as toothworts
(Dentaria), Rock cresses (Arabis), Winter and Water cresses (Barbarea), etc.
132
HOVANITZ
J. Res. Lepid.
Three broods. Adults in late April (Massachusetts).” "Not recorded s. of
the Catskill Mountains in New York.”
Clark (1951) says of virginiensis:
"Range :---One record, Frederick County northwest of Cross Junaion on
the Bloomery Road (Rank 698) about 200 feet east of the West Virginia
line, April 24, 1938.”
"Variation — This species varies considerably in the intensity of the
bordering of the veins on the under side of the hind wings. This bordering is
pale in ail the individuals from Virginia and adjacent West Virginia that we
have seen, some having the under side of the hind wings almost immaculate
white.”
"Occurrence: — Confined to rich deciduous woods and extremely local,
occurring in limited numbers at usually widely separated localities. The reduction
in the numbers of Pieris virginiensis and its present occurrence only in widely
separated localities are undoubtedly the result of deforestation, which has
greatly reduced the areas in which it is possible for it to maintain itself.”
"Pieris virginiensis keeps strictly to the woods.”
"Season: — One brood. This species appears shortly after the middle of
April and flies until about the middle of May.”
Clench (1958) writes of the same:
"Pieris virginiensis . . . common in woods . . . one brood: Late April to
mid-May. One . . . capture ... on 4 June, 1958 is surely a freak occurrence . .”
Scudder ( 1889) writes of the habits of oleracea:
"Food plants. This caterpillar feeds on various cultivated cruciferous plants,
such as turnip {Brassica rapa) , cabbage {B. oleracea), radish {Raphanus
sativa) , horseradish {Nasturtium armoracia) and mustard {S inapis ) . Mr.
Lintner considers that it prefers turnip to cabbage, for he once obtained fifty
eggs fom a turnip bed and could find none on adjacent cabbage plants of the
same age, and this has been my experience.” Hoy, on the other hand, thinks
it is found mostly on mustard. But it also occurs on some of our native plants,
such as Arabis drummondi (Couper) and A. perfoliata (Fletcher), and Fitch
says it occurs abundantly on the water cress {Barbarea vulgaris)."
"Life History. This butterfly, which appears to be generally triple brooded,
passes the winter in the chrysalis, the first brood has been seen as early as
April 18, but usually appears between April 27 and May 9, a week or there-
abouts after P. rapae. For several years Fitch observed it at East Greenwich,
N. Y. for the first time on May 8 and 9 and only once as early as May 6; in
northern localities it is sometimes as late as the third week in May. It usually
becomes abundant between the 10th and 15th of the month and continues
unti the end of the first week in June. The eggs are laid during the last half
of May and early in June, and hatch in from 5 to 10 days . .. . the second
brood . . . makes its advent during the last days of June or early July . . ., this
brood in oleracea seems never to be very abundant . . . and to maintain itself
for a comparatively short time; it becomes common by the end of the first
week in July and sometimes disappears . . . before the end of the month . . .
"The third brood appears as early as the last days of July but usually not before
August; it . . . lasts until early September . . . Occasional specimens, . . .
disclose butterflies late in September or early in Oaober.” "The species must
be triple brooded in the north.”
Edwards (1868-1872) wmts oi virginiensis:
"In the Kanawha district it replaces Oleracea which is yet unknown
there. It is not uncommon in the month of May, frequenting open woods
rather than gardens, and in this respect differing in habit from the allied
species. I have never met with it later than June, though Oleracea in the
(2):i24-i34, 196}
PIERIS NAPI RACES
133
Northern states is most abundant after that month and continues breeding
til the early autumn frosts.” "I have received specimens of Virginiensis from
Mr. Wm. Saunders of London, Canada and am informed by him that it is
there a rare insect.”
These data would tend to confirm the evidence derived from the
data of museum specimens that oleracea is indeed a muitivoltine race
which breeds from April all through the year until the Autumn frosts.
Our own personal experience on napi in California does not conflict
with this opinion. Here napi {venosa and castoria) are continuously
brooded when possible in relation to temperature and moisture condi-
tions. They go into diapause when conditions of the food plant are un-
favorable; this is usually in the summer when the food plant {Dentaria)
dries up. In fact much of the first winter generation does likewise as
Dentaria does not stay green long. This accounts for the summer or late
spring generations being less abundant than the winter or early spring
generations. A similar observation has been made of the generations of
napi in New England.
The food plant difference between the races oleracea and virginiensis
is important in only one way, namely in the restriction of habitat of
napi to a different ecological niche. Dentaria is a plant of the open
woods and partial shade. The flight period of virginiensis must be
adjusted to meet the proper conditions of moisture requirement for the
plant as well as for the degree of light availability in the deciduous
forest where early spring may allow more light in the forest than in
summer. The restriction of virginiensis to the "Transition” life zone
forest may be correlated with this light and temperature relationship.
The northern forest is cooler all summer long and the temperature needs
of oleracea can be met usually in the open areas as well as in the partial
shade of the forest. This would account for the different flight habits
of the two races. In California, napi ( venosa and castoria ) fly only in the
cool canyons where there is partial shade even though there is abundant
food available for them outside in the form of wild mustards. Instead
they are restricted to Dentaria of the shaded woods, while Pieris rapae
and P. protodice fly only a few yards away in the direct sunlight and
feed upon mustards.
The food plant of oleracea so often indicated in the literature is
winter or water cress {Barb area) which is apparently restricted to the
vicinity of strearqs. There is indication that in the Catskill Mountains
(New York) there may be a population of virginiensis on Dentaria
while sympatrically there are populations of oleracea on Barbarea. If
this is so, an excellent study could be developed to show the nature of the
racial or specific isolation of these two at a region where both maintain
reasonably distinct populations in close contact. The fact that oleracea
flies in April, virginiensis in May and oleracea again in June and July
indicates another additive isolation factor which would help to maintain
the species or racial distinctness.
By way of conclusion, the data presented show rather conclusively
that Pieris napi oleracea and Pieris napi virginiensis are not as com-
134
HOVANITZ
/, Res. Lepid,
pletely separated as one might wish "species” to be but that they are
not either fully racial or subspecific in the sense that their populations
blend into one another as by a continual gene flow. It is therefore a
matter of personal opinion whether they should be considered species or
races for nomenclatorial purposes. For these purposes it is impossible
to stand on the middle ground as a choice must be made. I choose to
use the racial designation for the reason that the groups are almost
completely allopatric, that is, the populations are geographically separ-
ated and do not occur in the same place with certainty without inter-
breeding and complete fusion. Also, intermediates do occur with high
frequency in intermediate geographical areas. There is work to be done
in this regard on the ground in important areas, such as, the Catskill
Mountains, He Perrot (Montreal) or any other areas where a potential
close unity of the two appears.
The past changes in the distribution of the species have been greatly
exaggerated, especially with regard to any competition between P, rapae
and P, napi. The habits of the two species are so distinct that no compe-
tition can reasonably exist. Past records of P, napi feeding on garden or
cultivated cruciferous plants are undoubtedly correct but these larvae
came from adults flying out of adjacent woods to lay their eggs on any
cruciferous plants available. With the destruction of the woods in the
New England states, the habitat for napi disappeared and then of
necessity napi disappeared. This was the situation for the low lands. In
the mountains, napi still exists as it did in the past.
REFERENCES
CLARK, A. H. 1951. The butterflies of Virginia. Smithsonian Misc. Col-
lections, 116(7) : 1-195.
CLENCH, HARRY K. 1958. The butterflies of Powdermill Nature Reserve.
Research Report #7. Powdermill Nature Reserve, Carnegie Museum,
Pittsburgh, Pa.
EDWARDS, W. H. 1868-1872. Butterflies of North America. Philadelphia.
The American Entomological Society.
FISKE, Wm. F. 1901. An annotated catalogue of the butterflies of New
Hampshire. New Hampshire College, Durham, N. H.
HOVANITZ, WILLIAM. 1949. Increased variability in populations follow-
ing natural hybridization. In Genetics, Paleontology and Evolution,
Princeton.
— — — — . 1962. The distribution of the species of the genus Pieris in
North America. /. Res. Lepid. 1 (1): 73-83.
KLOTS, A. B. 1935. On the life history of Pieris virginiensis Edws. /. N.Y.
Entom. Soc., 43: 139-142.
. 1951. A field guide to the butterflies of North America, East
of the Great Plains. Houghton Mifflin Co., Boston.
McHENRY, PADDY. 1962. The generic, specific and lower category names
of the Nearctic butterflies. Part 1. The Genus Pieris. /. Res. Lepid.
1(1): 63-71.
SCUDDER, S. H. 1889. The butterflies of the eastern United States and
Canada. Vols. I-III, Cambridge, Mass., author.
Journal of Research on the Lepidoptera 1(2) :135-156, 1963
1140 W. Orange Grove Ave., Arcadia, California, US, A.
© Copyright 1965
THE MALE GENITALIA OF SOME COLIAS SPECIES
BJORN PETERSEN
University, Lund, Sweden
In most families of Lepidoptera the genitalia show such
distinctive characteristics that it becomes, easy to separate even closely
related species. In the family Pieridae conditions are different. Pieris
mpi L., P. hfyoniae Ochs., and P. ergane Hbn. are not separable on
the basis of their male genitalia, nor are F. rapae L. and P. manni Mayer
(Lorkovic 1928, Drosihn 1933). Similar conditions are present in the
genus Colias.
Before going into details a general description of the male genitalia
of Colias will be given (cf. fig. 1). The distal part of the Vlllth tergum
is more or less slender, Kusnezov (1915) has called this part the super-
uncus, Warren (1950) named it the false uncus. At the base of the
superuncus the tergum is incompletely sclerotized laterally. The saccus
is rounded, sometimes pointed on its sides. The vinculum is long and
slender. The tegumen shows a narrow process dorsally, the pseudouncus
(Kusnezov 1915). The aedeagus has a long ventral arm.
According to Warren (Lc.) the claspers at their proximal end are
restricted to a blunt point, attached to the vinculum. The dorsal edge
is said to be drawn upwards, parallel to the vinculum, and the dorsal
terminal extremity attached to the tegumen. These statements do not
correspond to conditions found in Protocolias imperialis (PL I, fig. 1).
In this species the proximal part of the claspers is rather similar to that
of other Colias species. The distal part is protracted so that the clasper
obtains a shape more or less similar to that of many other Pierids.
It may therefore be concluded that the clasper -head with the terminal
tooth is the dorsal part of the clasper.
The short pseudouncus and the marked distal lobe of the clasper
give to P. impefialis a rather isolated position. In some characteristics it
comes closer to the genera Catopsilia and Anteos. In other respects,
however, such as the structure of the clasper head, P. imperialis is
similar to Colias, thereby showing the relationship between these genera
(cf.Klots 1929 a and b).
In the genus Colias the genitalia are rather variable within a species
or subspecies, as will be shown below. At the same time the differences
between some species are only slight. Therefore a biometrical approach
seemed necessary as a complement to the general descriptions of the
135
136
PETERSEN
J. Res. Lepid.
genitalia. The modes of measuring and the results are given first as
important for the full understanding of the descriptions.
The breadth of the superuncus. The edges of the superuncus in
dorsal view may a) converge distally b) run parallely or c) diverge
slightly from a narrow part near the base ( cf . fig. 2 ) . The breadth was
on diverging superunci measured at the narrowest basal point, on
others at the corresponding place.
Fig. 1. The male genitalia of a Colias. a superuncus b pseudouncus
c clasperhead d tooth of clasperhead e uncus
The size of a certain part of an individual may be influenced by its
general size. The breadth of the superuncus has therefore been correlated
with a measure of size: the wing length.
All species with genitalia similar to those of the three species
previously investigated: C. hecla, nastes and palaeno are included in
fig. 3. Here the breadth of the superuncus is plotted against the wing
length in a logarithmic scale. The populations investigated are parted in
two groups, the first with a slender superuncus, the second with a broad
one. The regression line of the populations with a slender superuncus
is:
log y 1= 0.75 log X + 0.68; (r = 0.72; 0.02 >£> 0.01).
The species along this regression line are both orange with a hecla-
pattern, yellow with a ^^c/^-pattern, and yellow with a nastes-patitm.
The species in fig. 3 with a broad superuncus are either orange with a
hecla-^RttQm or yellow with a nastes-pRttQm. These two types of species
can be grouped along two regression lines. Within both groups the
correlation is statistically significant.
t(2):i}5-i56, 196}
COLIAS GENITALIA
137
Orange species:
log y=: 0.80 log X + 0.77; (£ = 0.71; 0.05 > P > 0.02).
Yellow species:
log y 1.17 log X + 0.30; (r = 0.81; 0.05 > P > 0.02).
Fig. 2. Differently shaped superunci with semicircular, less sclerodzed areas
near the base., a convergent b parallel c divergent
The means of the two groups are, however, not significantly separated.
An analysis of covariance gives 0.1 > ^ > 0.05.
The data on which fig. 3 have been based are tabulated in Tables 1
and 2. Among the species shown in these tables C. nastes is of special
interest. In Scandinavia and in North America but also in Altai and
the Sayan Mountains the superuncus is slender. C. nastes of Siberia
and the Amur Province is intermediate between nastes of the Altai and
the Sayan Mountains and the closely related, allopatric C. montium from
S. Kansu. This species forms a transition to C. cocandica from Ferghana,
the Issykkul, and from the Tianshan. The superuncus-breadth of all
Asiatic individuals of nastes is plotted against wing length in fig. 4,
The populations are grouped along two regression lines, though the
correlation in none of the groups is statistically significant ( cf . Table 1 ) .
An analysis of covariance, however, shows that the means of the two
samples (shown in the figure by squares) are significantly different
(0.01 > P> 0.001).
In the yellow series of forms there is thus no sharp limit between
species with slender and broad superunci. Even within a single popula-
tion, that from the Amur Province — the variation is so great that it
includes superunci typical f(Jr nastes nastes and others typical for
cocandica (cf. fig. 4). In the orange series of forms conditions are
different as will be treated later. In fig. 5 the superuncus breadth of
arctic C. nastes and of its sibling in the Alps, C. phicomone, is plotted
against wing length. The differentiation has proceeded so far that
hardly any overlap is present.
The wing length and breadth of the superuncus in some other
Colias species, less similar to those treated above, are shown in Table 3.
In the three pairs at the top of the table the genitalia are so similar that
138 PETERSEN /• Lepid.
KOU J L I I ! I ( 1 I
1.30 1.32 1.34 l36 1.38 1.40 1.42 7.44 7.46 7.48
Fig. 3. Superuncus breadth of some Colias populations plotted against wing
length. Data from Tables 1 and 2.
O Population from N. America or northern Eurasia
□ Population from Central Asia or Central Europe
7\ C. palaeno and interior
rod 3hoNe=hecla patterns open mark=orange color
rod helow—nastes patterns filled mark=yellow color
no constant differences could be found. Warren (1950) states some
differences between C. australis and hyale, a statement which was not
confirmed by the specimens investigated.
Two of the species of Table 3, C. cunninghami and P. imperialis,
have an extremely broad superuncus. C. vautieri, which otherwise is
1(2): 135^156,
1.90 ^
.?«3 COLIAS GENITALIA
log superuncus breath
1.85
O
139
1.80
1.60
o +
log wing length
U. I -J_ — __
1.32 1.34 1.56 1.38 1.40 1.42 1.44
Fig. 4. Superanciis breadth of some Asiatic populations of C. nastes plotted
against wing length. The lower square represents the mean of the dots, the
upper square the mean of the remaining marks.
• Altai and Sayan Mts. A Transbaikal
O Siberia, Taiga zone + Amur Province
rather aberrant, C. erate, and myrmidone fit well into the group of
species with a broad superuncus, listed in Table 2. The remaining
four species have a more slender superuncus though C. hyale and
australis, like C. montium occupy a somewhat intermediate position.
The form of the superuncus in lateral view-. During copulation
PETERSEN
/. Res. Lepid,
140
Log superurunijS breadth
100^
95-
o
o
o
o
90-
85-
80-
O
O
O
O
o oo
o
o
o
o
o
o
o
75^
65-
60-
55-
126 1.50 tsz 134- 1.56
log wing Length
138 UO 142
Fig. 5 Superuncus breadth of C. phicomone (open circles) and C. nastes
(filled circles = Scandinavia, + North America) plotted against wing length.
(2):i}5~i56, 1963
COLIAS GENITALIA
.141
the superuncus together with the uncus is pressed against the body of
the female, just as the uncus in the genus Pieris according to Lorkovic
(1947). To make possible the bending down of the superuncus during
copulation a semicircular area laterally on the Vlllth tergit is less
sclerotized than the rest ( cf . fig. 2). This bending can be stated without
studying any copulating pairs. In some specimens the superuncus is
broken (cf. fig. 6), in some in addition kept between the two claspers.
Fig. 6. Colias intenor with superuncus broken during copulation.
The breadth of the superuncus of such specimens sometimes cannot
be measured without maceration. A broken superuncus was found in
all species where more than a few specimens were investigated, as
shown in Table 4.
Only in very few species the superuncus is a straight process
protruding from the Vlllth tergit. Also when unbroken the superuncus
usually forms a bow downwards. This bow is present, even in its most
pronounced form, in animals which have not copulated, as, for instance,
in all the thirty C. hecia specimens investigated which in 1952 were
caught at the beginning of the flying time on the northern side of
the Lake Tome Trask.
To obtain a quantitative estimation of the variation of the form of
the superuncus in lateral view, the angle of the distal part of the
superuncus to the dorsal edge of the Vlllth tergum was measured.
The result of this investigation is shown in Table 4. In general a
broad superuncus is only slightly bent downwards while a narrow is
bent more. The exceptions are rather few: C. interior, montium, and
phicomone.
142
PETERSEN
/. Res. Leptd.
Species
n
log X
M,
log y
r
P
b
A. Orange species
C, hecla, Scandinavia
51
1.331
1.70
-0.18
0.3-0. 2
-0.74
, Siberia, Dudinska
4
1.355
1.67
, Baffin, Isl.
2
1.347
1.79
, Greenland
3
1.362
1.74
C, hyperborea, N. Siberia
4
1,69
C. eurytheme, N. America
6
1.456
1.76
, slightly orange
Texas, Houston
6
1.313
1.61
B. Yellow species
a. hecla pattern
C. palaeno, Europe
41
1.411
1.71
0.18
0.3-0. 2
0.50
C. palaeno, Siberia
9
1.377
1.71
0.57
0. 2-0.1
1.73
C. interior
2
1.311
1.67
b. nastes pattern
C, nastes, Scandinavia
42
1.344
1.68
-0.09
0.6-0. 5
-0.24
, Siberia, Amur area
29
1.391
1.75
0.32
0.1-0.05
0.87
, Altai, Sayan Mts.
12
1.379
1.67
0.35
0.3-0. 2
0.86
, N, America
2
1.311
1.68
C. philodice, N. America
11
1.419
1.74
C. montium, S, Kansu
9
1.370
1.77
Table 1. Wing length (s<), in mm, and breadth of superuncus (y), in an arbitrary scale.
and their correlation in some Colias species with a slender superuncus. Logarithmic
scale.
C. interior is in this character well separated from its allopatric
sibling, C. palaeno, in having a rather straight but slender superuncus.
C. montium has a rather slender superuncus which is straight as is
that of cocandica. In this respect another of the southern species,
C, phicomone, is intermediate between nastes and cocandica. The
superunus of phicomone is, however, rounded and not more or less
pointed as it is in all other species with a slender superuncus except
C. interior, aurorina, and sagartia.
By means of the two characteristics, i.e. the breadth and the
form of the superuncus, it is possible to separate the two species
C. viluiensis Men. and hyperborea Gr. Gr. which live sympatrically in
N.E. Siberia. Of viluiensis 7 specimens from the Verchojansk area,
Lutsha near Yakutsk, Vilutsk and Vilui have been investigated. The
four specimens of hyperborea were from Sib. poL, the Lena Valley
I (2) ; I )5-i56, 7963
COLIAS GENITALIA
143
Species
n
”log X
M,
log y
Species
n
'^log X
M
log y
Orange species,
hecla-pattern
Yellow species,
nastes-pattern
C, chrysotheme
5
1.350
1.89
C. croceus, Europe
10
1.403
1.90
C. alpherahyi
1
1.436
2.02
, Asia
11
1.429
1.89
C. christophi
1
1.369
1.86
C. eogene
8
1.368
1.87
C. heos
12
1.474
1.99
C. cocandica
2
1.326
1.85
C. remanovi
2
1.446
1.91
C. phicomone
24
1.374
1.88
C. standingeri
6
1.395
1.83
C. sieversi
1
1.405
1.93
C. thisoa
2
1.417
1.90
C. sifanica
1
1.326
1.90
C. wiscotti
9
1.431
1.93
C. viluensis
7
—
1.90
Table 2. Wing length (x), in mm, and breadth of superuncus (y), in an arbitrary
scale, ih some Colias species with a broad superuncus. Logarithmic scale.
and Sredne Kolymsk.. The latter specimens all have a slender and
strongly bent superuncus of the hecla-type, while all specimens of
viluiensis have a straighter and broader superuncus. There is no overlap
in any of the two characters; in the "angle-character” the gap is very
wide.
In Colias nastes from Asia there Is no similar correlation between
the breadth and the shape of the superuncus. Among the 41 specimens
investigated the coefficient of correlation is -j- 0.093 which is far below
significance ( 0.8 > P > 0.7 ) .
Conditions are thus quite different in northern Asia as regards the
nastes- and hecla-stn^s of forms. In the nastes-SQnts dines including
characters of the genitalia reach the Amur area, Transbaikal, the Sayan
Mts, and the Altai, and forms intermediate between arctic and Central
Asiatic ones are present in North Western China and in the mountains
of Central Europe. In the orange series of forms an overlap of a northern
and a southern species is present between 65° -68° n. latitude and no
intermediates have yet been found between these species.
Number of teeth near apex of aedeagus. In many groups of insects
the number, shape, and position of the aedeagal teeth serve as good
evidence to distinguish species. In the Colias species all the teeth are
small, of a rather similar shape, situated near the apex. Variation in
number is strong even within subspecies when compared with the
differences occurring between species. Thus this character is without
signifiance for the determination of individuals. A closer investigation
of a material belonging to the group of species with a slender super-
144
PETERSEN
/. Res. Lcpid.
C. aurorina
n
M,
log X
M,
log y
6
1,449
1.80
C, sagartla
5
1.437
1.76
C. australis
8
1.384
1.74
C. hyale
27
1.373
1.79
C . erate
13
1.387
1.94
C, myrmidone
7
1.425
1.95
C. cunninghami
2
1.367
2,01
C. vautieri
2
1.327
1.03
P. imperialis
1
1.356
2.05
Table 3. Wing length (x), in mm, and breadth of superuncus (y),
in an arbitrary scale, in some Colias species.
uncus, however, revealed some slight specific differences in the average
number of teeth (cf. Table 5). The results of this investigation are;
C. nastes has on an average fewer teeth than C. hecla from the
same locality (Scandinavian material t = 2.45; 0.02 > P > 0.01,
American material not differing significantly. There is a certain tendency
of parallelism, both species having a lower number of teeth in Scandi-
navia than in North America. This tendency is not significant in any
of the species, however. C. palaeno and interior both have a low number
of teeth, while the number is fairly high in C. meadi, philodice, and
eurytheme.
SPECIAL PART
The genus Colias may be parted in two genera: Protocolias (type
imperialis Btlr.) and Colias, the latter in turn in two subgenera
Mesocolias (type vautieri Guer.) and Colias. The descriptions of the
genera, the subgenera and their various species may be given most
easily in the form of a key.
1. Small or medium-sized, orange butterflies with broad (0.40-0.45
mm) or very broad (0.5 mm), straight superunci. Superficial scales
— if present— -yellow and black, broad and flattened in the distal
part. Apex of aedeogus without teeth.
Genus Protocolias and subgenus Mesocolias 2.
r. Small to big, yellow, greenish or orange species with broad to
slender (0.32 mm) superunci. Superficial scales in the black mar-
gin—if present— yellow, pointed, hairlike or broader. A number
of teeth at apex of aedeagus
Subgenus Colias 3.
i(2):i}5-i56, 196}
COLIAS GENITALIA
145
2. Superuncus very broad and triangular. Pseudouncus short. Clasper-
head with elongated tooth. Middle part of the clasper with a
marked prong. Ventral arm of aedeagus broader in the distal end.
Black and yellow suoerficial scales present.
Pfotocolias imperialis Btlr. (PI. 1:1)
2’. Superuncus broad. Tooth of clasperhead slightly bent upwards.
Ventral lobe (v.l.) of inner side of the clasper (Klots 1929 a)
more marked than in any other species of the genus. Distal part of
ventral arm of aedeagus showing charaaeristic shape. Superficial
scales absent.
Colias (Mesocolias) vautieri Guh. (PI. 1:2,3)
2”. Superuncus very broad, genitalia small, claspers short ( 1.0-1. 1 mm)
but broad (0.6 mm). Distal part of ventral arm of aedeagus not
broader than the proximal one. Superficial scales absent.
Colias {Mesocolias) cunninghami 'Qth. (pi. 1:4).
3. Claspers caudally with a marked prong directed medially. Super-
uncus straight and fairly slender.
C. hyale L. (pi. 1:5) and australis Ver.
3’. Claspers without any marked prong ........4.
4. Claspers pointed in the middle part of the caudal edge
(most easily seen from behind) ........5.
4’. Claspers not pointed in the middle part of the caudal edge 6.
5. Superuncus straight, on an average 0.40-0.45 mm broad
C. erate Esp., C. myrmidone Esp. (PI. 1:6),
5’. Superuncus usually markedly bent downwards, 0.30-0.35 mm broad.
C, sagartia Led. ( PI. 1 : 7 ) C. aurorina H. Sch. (PI. 1:8).
6. Superuncus on an average less than 0.32 mm broad (cf. text fig. 3)
usually strongly bent downwards (cf. Table 4) .........7.
6’. Superuncus on an average broader than 0.35 mm, usually
straight 13.
7. Superuncus only slightly bent downwards ..........8,
7’. Superuncus strongly bent downwards 9.
8. Superuncus in caudal view blunt-ended as in species with a broad
superuncus. Inner side of -clasper of hecla-type (cf. PI. 11:34)
C. montium Oberth. (PI. 1:9, 10).
8’. Superuncus in caudal view pointed. Inner side of clasper with well
developed ventral lobe and ridged from its dorsal edge dorso-
caudally towards the caudal part of the clasper (Fig. 14, 15)
C. interior Scudd. (PI. I: 11-13, II: 14, 15),
9. Ridge from dorsal edge of ventral lobe towards caudal part of the
clasper present as in C. interior. Ventral lobe less well developed
than in C. interior (cf. Fig. 16). Caudal edge of clasper often
strongly bent inwards ( most easily seen in caudal view ) .
C. palaeno L. (PI. 11:16-25).
C. Christina (PI. 11:26-28).
C. palaeno: is most variable in all charaaers investigated except
the breadth of superuncus (cf. the figures). One specimen from
Abisko (42 specimens studied) completely lacks the ridge on the
inner side of the clasper. Hence it cannot with certainty be separ-
ated from C. nastes or hecla. The limited material of C. Christina
appeared rather close to some specimens of palaeno, though it
might be possible to separate the two species after examining a
greater material.
9’. The caudal edge of the valva less strongly bent inwards ventrally.
Inner side of clasper reminding of hecla-type (cf. fig. 34) ........10.
10. Clasoer (in lateral view) broadest rather ventrally 11.
10’. Clasper broadest in the middle part 12.
11. Aedeagus on an average with a higher number of teeth at the distal
end ........................................C hecla Lef. (PI. 11:29-39).
146
PETERSEN
/. Res. Lcpid.
Frequency of
superuncus
broken
7/50
1 14/56 1
1 14/56 1
14/56
1/12
4/25
Ipdl 1
1 1
2/9 1
2/12
<D
DO
C
<
82
s
s
cr>
ID
s
s
<T>
CD
s
O
to
r-
.It
CO
El
CM
=t-
o
-
95-104 1
CM
-
c.
-
*— J
if
-
-
CO
CO
-
J-
-
-
3
-
-
*— 1
-
•— J
-
CD
-
f—{
CO
d-
S
-
-
-
-
-
-
ID
-
-
it
*— 1
CM
-
-
-
3
m
a-
-
-
rH
fH
o
c
-
CM
#— J
-
-
CO
-
25-341
tD
-
-
CN
1
•H
CM
•— 1
it
•H
Species,
Population
Popu of Table 1.
C. hecla, Scand. Mts.
" , N. America
C. hyperborea
C. meadi
j C. eurytheme |
C. palaeno, Scand. Mts.
j ” , Europe |
1 " , Asia
" , N, America
C. interior [christina]
m
■p
'O
c
o
CD
(0
«D
CO
m
c
m
CO
<
" , N. America
1 C. philodice
1 C. beryl la
Pop. of Table 2.
C. chrysotheme
i
C. croceus, Europe
0}
c
0)
DO
0
o
IT. Aedeagus on an average with a lower number of teeth. Very
similar to C. hecla .....C, nastes Bdv. (PI. III:35"38).
12. Bigger, superuncus broader ......C. eurytheme Bdv. (PL 111:39, 40).
C. philodice Gdt. (PI. III:4l).
12’. Smaller, superuncus slenderer. ....C. meadi Edw. (PL 111:42, 43).
13. Superuncus bent downwards C. phicomone Esp. (PL 111:44).
14. Superuncus diverging, very broad near the distal end. Clasper (in
lateral view) broadest ventrally C. croceus Fourcr. (PL 111:45).
14’. Superuncus often with parallel or converging edges or clasper
broadest in the middle part 15.
15. Clasperhead more strongly marked, tooth strongly developed. Ven-
tral arm of aedeagus broad in the distal part.
C. wiscotti Stgr. (PL 111:46, 47)
C. marcopolo Gr. Grsh. probably also belongs here. No material
has been available for investigation. Professor Sheljuzhko informs
I (2); 1)5-156, 196)
COLIAS GENITALIA
147
2/10 I
CO
1
4/30
2/15
0
0
0
fO
0
CD
CM
0
fO
0
CO
CD
to
0
0
s
CO
CO
o
.1^
o
CO
o
CM
o
zS-
-
e.
CM
f—h
-
-
c
-
CM
-
CO
CM
-
eH
iH
-
-
-
-
CM
CO
-
CO
-
-
OJ
CO
CM
rH
t—i
CM
-
-
-
-
j C. heos 1
C. romanovi
C. staudingeri
C. thisoa
C. wiscotti
1 C, vlluensis |
j C, alpherakyi j
C. christophi
j C. cocandica
i
B
0
0
U
j C, sieversi |
C. sifanica
Pop. of Table 3.
C. aurorina
C, sagartia
j C« australis |
C. hyale
C. erate
j C* rayrmidone j
C. cunninghami
! C. vautieri |
j P. imperialis j
that Avinov and Kusnezov both place marcopolo together with
wiscotti, the latter as a subspecies. According to Sheljuzhko both
forms fly together in the Pamir area and are probably different
species in spite of a great similarity in the genitalia.
15’. Clasper head less strongly marked 16.
16. A number of species remain which due to lack of sufficient
material and/or great similarity could not be separated from each
other :
Yellow species with nastes-p2XtQtn\ C. dpherakyi Stgr. (PI.
111:48), C. chfistophi Gr.-Grsh. (PL IV:49), C, cocandica Erscii.
(PI. IV: 50) (Specimen figured with triangular saccus; in a second
specimen investigated the saccus is rounded as in all other species),
C, sieversi Gr.-Grsh. (PI. IV:51), C. sifanica Gr. Grsh. (Pi.
IV: 52). These Central Asiatic species together with C. nastes,
phicomone, montium and others form one or — most probably —
two polytypic species.
148
PETERSEN
/. Res. Lepid.
n
2
3
4
5
6
7
0
9
10
11
12
M
C. hecla, N. Am.
8
5
1
2
6.6
*' , Greenl.
3
1
1
1
5.0
" , Scand.
30
2
7
9
8
4
6.2
C. meadi
10
4
1
3
1
1
7.6
C. eurytheme
12
3
3
5
1
6.4
C. palaeno, N. Am.
10
1
1
5
2
1
5.2
'* , Asia
8
1
1
6
5.5
" , Estonia
8
1
3
2
1
1
6.1
*' , Scand.
21
1
2
7
2
3
1
3
1
1
5.4
C. interior
11
3
4
2
2
5.3
C, Christina
1
1
6
C, nastes, N. Am.
6
1
4
1
6.2
*' , Asia
5
1
3
1
6.0
” , Scand.
23
1
4
8
5
5
5.4
C. philodice
5
1
2
1
1
7.0
Table 5, Number of teeth near the apex of the aedeagus of Colias species with a slender
superuncus. Orange species with hecla-pSLttern: C, chrysotheme Esp. (PI.
IV:53), C, eogene Fldr, (PI. IV; 54, 55), C. staudingeri Alph.
(PI. IV: 56), C. thisoa Men. (PI. IV:57), C. viluiensis Men.,
and C. heos (PI. IV:58, 59). As in the previous group it is un-
certain whether all these forms deserve each a specific status.
Orange species with «^j/^j-pattern : C. romanovi Gr. Grsh.
(PL IV:60).
This enumeration of Colias species is not complete.
DISCUSSION
The key given above does not show the phylogenetic relationships.
These are, due to extensive introgressive hybridization, very difficult
to find out. Suppose species A is closest related to the allopatric species
B which has given rise to species C sympatric to A. Due to the
introgressions A C or C A it is then possible that A and C
are more similar in several characters than A-B or B-C. However, some
characteristics, as for instance those on which the isolation between A
and C depend, are likely to be of maximum difference in the case A-C.
Among the Co/m-species investigated there exists a number of
pairs of species within which the male genitalia are very similar:
C. eurytheme-philodice; C. hecla-nastes; C. aufofina-sagartia; C. myrmi-
done-erate; C. hyale-australis and probably also C. pdaeno-christina and
C. wiscotti-mar copolo. Introgression has among these been established
between C. eurytheme-philodice (Hovanitz 1949 a, b) and between
C. hecla and nastes. Hybridization has been suggested between C. sugar-
tia and aurofina (Lederer 1941). It is possible that pairs of a similar
kind are present in Central Asia among the species figured from number
(2);i}5-i56, 1963
COLIAS GENITALIA
149
48 to number 58. Lederer (1941) mentions, among other suspected
hybrids, specimens which have been supposed to be hybrids between
C. eogene and cocandica. A number of species with different genitalia
have been seen in copula ( Lederer l.c. ) : hyde x myrmidone, hyde x
croceus, hyde x erate, croceus x erate, and hyde x phicomone. The last
cross gave rise to some larvae which died before the pupation. All
these species have markedly different genitalia and are probably not
very closely related. It therefore seems uncertain whether hybridization
in these cases can give rise to introgression. If introgression occurs it
must be possible to observe this fact also on the . genitalia.
In all the pairs of sibling species first mentioned {eurytheme-
philodice etc.), except hyde-austrdis and pdaeno-chfistina, colors, and
in most cases also patterns, are different. The first species is usually
orange with a heda-p2XXQm, the second yellow with a nastes-^2XX^iVi.
Only C. sagarda has in addition a blue pigment which is present in
some individuals of its orange sibling aurorina.
The common occurrence of differences in color and pattern between
sibling species suggests that these colors are integrating part of the
isolating mechanisms within the pairs. Sexual isolation of this kind has
in the pair Pieris napPbryoniae been established by Petersen, Tornblom
and Bodin (1952). Males of both species are attracted by the white
color of the P. napi female. The yellow female of bryoniae attracts the
males solely by means of movements and odors. No releasing effect of
any of the types of pattern was obtained neither in these pierids nor
in similar experiments with the Silverwashed Fritillary {A. paphia L.)
(Magnus 1954). Different color but probably not different pattern may
therefore play a role for the sexual isolation between Colias species.
The geographical distribution of some subdivisions of the genus
Colias may also be discussed. Protocolias and Mesocolias are entirely
South American. The species of the subgenus Colias with a broad
superuncus are all Palearctic, one group having penetrated even into
the Ethiopian region. Colias with a slender superuncus are mainly
distributed in North America and the northern Palearctic, only a few
living further south in the latter region.
The distribution of the groups is to a great extent certainly the ..
result of an evolution within different areas. It does not seem established
that palearctic Colias have evolved from South American forms. They
may as well have developed from primitive Colias in some other part
of the world where they are now extinct.
The evolution of the superuncus. Among the two characteristics
of the Colias genitalia, the superuncus and the pseudouncus, the latter
is present in the genus Anteos as well as in Catopsilia and Colias (Klots
1929 a, b, Drosihn 1933). The superuncus, on the other hand, is very
small and triangular in the Anteos species investigated {menippe,
clofinde, Plate IV, figs. 61, 62) covering only the pseudouncus and the
basal part of the uncus. A similar, though bigger superuncus is found
in Protocolias imperialis and the genus Phoebis (Drosihn 1933). In
150
PETERSEN
/. Res. Lepid.
the latter genus no pseudouncus is present. The Catopsilia and Aphrissa
species have broad, straight and diverging superunci (cf. Plate IV,
figs. 63, 64 and Drosihn 1933), rather similar to those of some Colias
species. The triangular shape of the superuncus is probably primitive, as
this shape is present in Anteos where the superuncus is comparatively
small.
The superuncus has in the genus Colias (and probably also in
Catopsilia and Aphrissa) taken over the function of the uncus of many
other Lepidoptera (cf. Lorkovic 1947) to assist medially and dorsally
in holding the female body during the copulation. The superuncus
and the pseudouncus probably developed to support the uncus dorsally.
As the superuncus became larger, it was placed directly against the
female body, and thus the uncus instead changed to support the super-
uncus.
It has been suggested, for instance by Verity (1947), that Colias
have developed from the Old World Catopsilias. The presence of the
very primitive Protocolias with a triangular superuncus, a short pseudo-
uncus, and the middle part of the clasper with a prong, makes this
assumption rather unlikely. The triangular superuncus may be con-
sidered as a very pronouncedly convergent one ( cf . fig. 2 ) . The more
or less parallel or divergent superunci of Colias, Catopsilia and Aphrissa
may have evolved by means of parallel evolution towards more uncus-
like conditions as discussed below in the case of Colias.
The Catopsilias and most Colias species have broad, straight, and
unpointed superunci, characteristics which therefore may be considered
primitive compared with those of C. hecla, nastes and palaeno. A
similiar result is arrived at, if the problem is approached from another
direction. As already mentioned the superuncus has in the genus Colias
taken over the function of the uncus. The latter has had its function
during so much longer a time that it may give a certain indication of
what shape is most apt to give an optimal function. The uncus is
slender, pointed, and rather straight. In the two first of these character-
istics the uncus corresponds to the superuncus of the hecla-nastes-
palaeno-x^^t. Only it is even more pronounced then. The shape of the
uncus on the other hand, is straighter in lateral view. The basal part of
the superuncus is, however, situated more dorsally than the same part
of the uncus ( cf . textfig. 13). Only if the superuncus is bent down-
wards, it can be placed against the female body in the same place as
the uncus of other Pierids.
The evolution of the superuncus thus seems to have converged
with that of the uncus. There is, however, still a marked difference in
shape between these two organs and for different reasons it is not likely
that the convergence will become complete ever. As already mentioned
the situation of the two organs is different. The pressure of the super-
uncus is supported by that of the uncus and probably also by that of
the pseudouncus, and finally the claspers are in the genus Colias not
built as in other genera.
(2):i)5-i56, 1963
COLIAS GENITALIA
151
SUMMARY
The male genitalia of a number of Colias species have been de-
scribed. The genus is divided into one new genus and two subgenera
of the genus Colias: The South American genus Protocolias (type
imperialis Btlr.) and subgenus Mesocolias (type vautieri Guer.) and
the mainly Holarctic subgenus Colias. The latter can be divided into a
palearctic group with a broad straight superuncus and a nearctic and
northern palearctic group with a slender superujicus which is bent
downwards. Transitions between these two groups exist within the
supraspedes C. nastes.
The evolution of the superuncus from a small beginning, as at
present in the genus Anteos, via the broad superund of the Catopsilias
and some Colias into a slender superuncus has been discussed.
Several pairs of Colias species with identical or very similar geni-
talia exist. It is suggested that the similarity is combined with intro-
gression. Several of the pairs include species of different colors. These
colors may serve to isolate the species sexually from each other.
REFERENCES
DROSHIN, J- 1933. Uber Art-und Rassenunterschiede der mannlichen
Kopulationsapparate von Pieriden, 1-135.. Stuttgart.
HOVANITZ, W. 1949a. Increased variability in populations following
natural hybridization. In Jepseo, et al. Genetics, Paleontology, and
Evolution. 339-355. Princeton.
HOVANITZ, W. 1949b. Interspecific matings between Colias eurytheme
and Colias philodice in wild populations. Evolution. 3: 170-173.
KLOTS, A. B. 1929a. The genus Anteos Hubner ( Lepidoptera, Pieridae).
Bull. Brooklyn Ent. Soc. 24: 134-214.
^ 1929b. The generic status of Catopsilia Hubner and
Phoebis Hubner, with a discussion of the relationships of the species and
the homologies of the male genitalia (Lepidoptera, Pieridae). Bull.
Brooklyn Ent. Soc. 24: 203-214.
KUSNEZOV, N. J. 1915. Insectes Lepidopteres I in Faune de la Russie
1-337. Petrograd.
LEDERER, G. 1941. Die Naturgeschichte der Tagfalter. Stuttgart.
LORKOVIC, Z. 1928. Analyse des Speziesbegriffes und der Variabilitat der
Spezies auf Grund von Uotersuchungen einiger Lepidopteren. Glasn.
Soc. Scient. Nat. Croat. 38: 1-64,
LORKOVIC, Z, 1947. Modes artifidels d’accouplement des papillons.
Glasn. Biol. Sekc. Ser. II/B 1 :86-98.
MAGNUS, D. 1954. Experimentelle Untersuchungen am Kaisermantel zur
Analyse optischer Auslosungsreize. Deutscher Entomologentag in Ham-
burg 30. Juli bis 3. August 1953: 58-75.
PETERSEN, B., O. TORNBLOM, and N. -O. BODIN. 1952. Verhaltens-
studien am Rapsweissling und Bergweissling (Pieris napi L. und Pieris
bryoniae Ochs.) Behaviour. 4: 67-84.
WARREN, B. C. S, 1950. Speciation in the genus Colias: with special
reference to C. hyale and C. australis. Lambillionea. 50: 90-98.
152
PETERSEN
/. Res. Lepid.
PLATES
a. lateral view
b. superuncus, dorsal view
c. caudal view
d. aedeagus, ventral or dorsal view
e. ” , distal end in lateral view
f. clasper from inner side
g. saccus in ventral view
u. uncus
v.l. ventral lobe of the inner side of the clasper (Klots 1929 a)
PLATE I.
Fig. 1. P. imperialis (wrongly labelled
Honolulu)
Fig.
2.
C. vautieri, Ensenada
Fig.
3.
C. vautieri, Ensenada
Fig.
4.
C. ctinninghami, Junin, Peru
Fig.
5.
C. hyale, Oland, Sweden
Fig.
6.
C. myrmidone , Germany
Fig.
7.
C. sagartia, N.E. Persia
Fig.
8.
C. aiirorina, Armenia
Fig.
9.
C. montium, Tatsienlou, Tibet
F.g.
10.
C. moiitium, S. Kansu, China
Fig.
11.
C. interior, Montreal, Canada
Fig.
12,
13.
C. interior, Alaska Highway, mil
126, Beatton R. area, B.C.
PLATE
II.
Fig.
14.
C. interior, Alaska Highway, mil
90, Beatton R. area, B.C.
Fig.
15.
C. interior, Alaska Highway, mil
90, Beatton R. area, B.C.
Fig.
16.
C. palaeno, Alaska Highway, mil
450, Toal River, B.C.
Fig.
17.
C. palaeno, Bjurfors, Sweden
Fig.
18.
C. palaeno, Sweden
Fig.
19,
20.
C. palaeno, Smaland, Sweden
Fig.
21.
C. palaeno, Abisko, Sweden
Fig.
22,
23.
C. palaeno, 10 miles South
Burwash Landing, Y.T.
Fig.
24,
25.
C. palaeno, Sweden
Fig.
26.
C. Christina, America borealis
Fig.
27.
C. Christina, Rocky Mts.
Fig.
28.
C. Christina, Am. bor.
Fig.
29.
C. hecla. Nr. Haines Junction,
Y.T., up Summit Cr.
6000’-7000’ el.
Fig.
30.
C. hecla. Bog. nr. Johnson’s
Crossing, Y. T.
Fig.
31.
C. hecla, Dudinska, Siberia
Fig.
32.
C. hecla, Sweden
Fig.
33.
C. hecla, Kvikkjokk, Sweden
Fig. 34. C. hecla, 20 miles South
Burwash Landing, Y. T.
PLATE III.
Fig.
35.
C. nastes. Nr Haines Junction,
Y. T., up Summit Creek,
6000’-7000’ el.
Fig.
36.
C. nastes, Mt. Atabaska, Jasper
N.P., Alberta, 7000’-8000’ el.
Fig.
37,
38.
C. nastes, Kvikkjokk, Swedish
Lapland
Fig.
39.
C. eurythemc, Texas
Fig.
40.
C. eury theme , Minnesota, U.S.A.
Fig.
41.
C. philodice, Amer. bor.
Fig.
42,
43.
C. meadi. Bow Pass, Jasper N. P.,
Alberta
Fig.
44.
C. phicomone, Alps
Fig.
45.
C. croceus, Tirol, Austria
Fig.
46.
C. wiscotti, Turkestan
Fig.
47.
C. wiscotti separata, Turkestan
Fig.
48.
C. alpherakyi , Turkestan
PLATE
IV.
Fig.
49.
C. christophi, Turkestan
Fig.
50.
C. cocatidica, Turkestan
Fig.
51.
C. sieversi, Turkestan
Fig.
52.
C. sifanica
Fig.
53.
C. chrysothemc
Fig.
54.
C. eogene, Kisil Fast area
Fig.
55.
C. eogene
Fig.
56.
C. staudingeri, Tian Shan,
Fu-Shu-Shan
Fig.
57.
C. thisoa
Fig.
58.
C. heos, N. Mongolia, long. 100°-
110°, lat. 45°-50°
Fig.
59.
C. heos vespera, S. Kansu, China
Fig.
60.
C. romanovi, Turkestan
Fig.
61.
Anteos menippe, Mattogrosso
Fig.
62.
A. clorinde, Valles, Mex.
Fig.
63.
Catopsilia florella, Syria
Fig.
64.
C. grandidieri, Madagascar
i(2):t35-i56, 196} COLIAS GENITALIA 153
PLATE I
PETERSEN
PLATE II
COLIAS GENITALIA
PLATE III
156
PETERSEN
/. Res. Lepsd.
PLATE IV
6fb\
Journal of Research on the Lepidoptera 1(2) :157-162, 1963
1140 W'. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 1965
P
THE EFFECT OF HYBRIDIZATION OF HOST-PLANT
STRAINS ON GROWTH RATE AND
MORTALITY OF PIERIS RAPAE'
WILLIAM HOVANITZ and VINCENT C. S. CHANG
California Arboretum Foundation, Inc., Arcadia,
Los Angeles State College and
University of California, Riverside
It has been previously shown (Hovanitz and Chang, 1962)
that larvae of strains of Rieris rapae differ in their mortality rate, growth
rate and final size dependent upon the plant which is supplied them for
food. Slightly greater viability, greater rate of growth and increased
size are characteristic results of feeding the larvae on kale (Brassica
oleracea var. acephala) as compared with mustard (Brassica nigra),
even though both plants are utilized by the species in the wild.
It is the purpose of this paper to report on the results of an experi-
ment designed to compare these physiological factors in strains bred
for several generations on each plant 'and then hybridized.
THE STRAINS
The strains of Rieris rapae used in these experiments were derived
originally from two sources, both in the Los Angeles Basin of southern
California. The first, here designated the kale strain, originated with
wild females obtained in a cabbage field (Brassica oleracea var. capitata)
in a truck crop growing area in western Orange County, near Hunting-
ton Beach. The other, here designated as the mustard strain, originated
in the fields of the Los Angeles State and County Arboretum, where in
the spring time Rieris rapae may be found in conjunction with black
mustard (Brassica nigra). These have probably had no recent contact
with cabbage since there are none grown commercially within nine
miles of the area. Before testing in the experiments here described, the
kale strain had passed through more than ten generations in the labora-
tory on kale and the mustard strain had passed through more than six
generations in the laboratory on mustard.
'Aided by a grant from the National Science Foundation, Washington, D. C.
157
158
HOVANITZ AND CHANG
/. Res. Lepid.
THE COMPARATIVE TESTS
Six comparative tests were run contemporaneously, consisting of
twenty larvae each. These larvae were fed in petri dishes by daily
changes of food, keeping each larvae well isolated from the others. It
was especially important while they were small that the larvae were
placed directly on top of the leaf of the plant. Of the six tests, two
were of the parent strains; one of these was derived from kale and fed
on kale during the experiment, while the other was derived from
mustard and fed on mustard during the experiment. Four further sets
were Fi larvae from these two parent strains derived as follows:
One set consisted of Fi larvae obtained by crossing a kale-strain
female with a mustard-strain male and then by feeding the ensuing
larvae on the food-plant of the mother ( kale ) .
A second set consisted of larvae obtained by crossing a mustard-
strain female with a kale-strain male and then feeding the ensuing Fi
larvae on the food plant of the mother ( mustard ) .
A third set consisted of larvae obtained as in the manner of the
first F 1 set above, but fed on the food plant of the father ( mustard ) .
A fourth set consisted of larvae obtained in the manner of the second
F 1 set above, but fed on the food plant of the father ( kale ) .
MORTAUTY
The mortality figures for the larvae were low in all the strains
( table 1 ) . The kale-strain parents had a mortality of ten percent, which
compared well with the data previously obtained ( Hovanitz and
Chang, 1962). The mustard-strain parents had a mortality of thirty
percent which also compared well with the previously recorded data.
The reciprocal Fi’s behaved differently. When the kale-strain con-
tributed the female parent and the mustard-strain contributed the male
parent, the mortality of the Fi larvae was twenty percent whether kale
or mustard was used as food. On the other hand, when the mustard-
strain contributed the female parent and the kale-strain contributed the
male parent, the larvae had only a ten percent mortality when used on
kale and zero percent mortality when bred on mustard. It is doubtful
that these slight differences are significant for the N value of 20, though
they may represent a real difference. It is possible that the hybrids show
a greater viability as a result of their heterozygosity. Both parental
strains have been considerably inbred during their maintenance in the
laboratory. No reason can be given at this moment for the greater
viability of the mustard female F/s (10 and 0 percent) as compared
with the reciprocal kale female F/s (20 percent each).
i(2):t5y-i62, 196}
PIERIS HYBRIDIZATION
159
LENGTH OF LARVAL PERIOD
The minimum length of life for the larvae from the egg to the
adult for both of the parental strains was twenty-five days under the
conditions of the experiment (table 1) . All four of the hybrid F, strains
had a shorter larval length of life indicating a greater activity than the
inbred parental strains. The rate was speeded up to twenty-two days for
the Fi strains in which the female parent was derived from kale stock.
Type
Strain
No.
No.
No.
and %
Minimum days
from egg to pupae
Kale-bred
5
20
2
10%
25
Parents
Mustard-bred
6
20
6
30%
25
u
Kale-bred
2
20
4
20%
22
Kale ^ X Mustard ^
Mustard-bred
1
20
4
20%
22
u
Kale-bred
4
20
2
10%
21
Mustard ^ x Kale 6^
Mustard-bred
3
20
0
0
23
Table 1, The comparative mortality and length of larval growth
period of various strains of Pieris rapae.
For the Fi stock in which the female parent was derived from mustard,
the larvae bred on kale were speeded up to twenty-one days and those
bred on mustard to twenty-three days. These data definitely indicate a
significantly greater activity of the hybrid strains than the parental.
However, there is no indication of any maternal effect on the inheritance
of food-plant preference since the reciprocal crosses indicated the same
general effects on mortality and growth rate whichever strain con-
tributed the female parent.
GROWTH RATE AND MAXIMUM SIZE
The growth rate of the six sets of larvae was compared by measur-
ing ten of the living larvae daily from the time of hatching from the
egg to the time of pupation. The average of these ten measurements
was plotted daily and curves drawn to illustrate their size increases
(fig. 1). Differentiation between the curves occurred within the first
few days but the first five days have been omitted in fig. 1, owing to
the small size of the larvae. On the fifth day, both parental strains had
the smallest larvae while the two F, strains, in which the kale-strain
female parent was used, had the largest larvae. The other two Fi strains,
in which the mustard- strain female parent was used, were intermediate.
During the entire growth period, nearly the same relationships held^
with the exception of the kale-bred Fi with a mustard-strain female
160
HOVANITZ AND CHANG
/. Res. Lepid.
Fig. 1. Growth rates of larvae of six strains of Pieris rapae on kale and
mustard. Vertical scale, length in mm.; horizontal scale, days after
hatching from egg to pupation.
(2):i5j~i62f 1^63
PIERIS HYBRIDIZATION
161
parent which for a time was faster than aM the others. Of the two
parental groups, the mustard -bred larvae were larger than the kale-bred
at five days, but the kale-bred reached a maximum ahead of and larger
than the mustard-bred. Of the three groups on kale as compared with
the three on mustard, the kale-bred group in each case reached a maxi-
mum size larger than or earlier than the comparable group on mustard.
Of the parental as compared with the Fi groups, the larvae of both
F] hybrid groups consistently exceeded the larvae of the parental
groups from the beginning to the end.
The largest maximum size was attained by the F\ group in which
the female parent was derived from a kale-strain and in which the
larvae were kale-bred. On the other hand, the second largest maximum
size was attained by the larvae of the F] group bred on mustard but
which had a mustard-strain mother.
It may be observed from the curves that at the periods of molting,
sizes of the larvae are reduced. These periods are shown by the arrows
(fig. 1). Because these reduction periods are delayed for the slower
larvae, the older the larvae, the more difficult it is to compare the six
groups on any one day.
SUMMARY AND CONCLUSIONS
1. Crosses were made between strains bred for many generations
on kale and mustard.
2. The effect of hybridization between these host-selection strains
has been to speed up development, to bring about increased sizes to
the larvae, and to reduce mortality. These effects are probably due to
the genetic effects of "hybrid vigor”, in which the deleterious effects of
inbreeding within the host-selected strains have been reduced by out-
crossing.
3. These effects are greatest for the kale-strain than for the mustard-
strain. This would tend to confirm our previous findings that- kale is
preferred as a food over mustard, even when mustard is utilized as the
food for many generations.
4. The increased viability, increased growth rate and increased size
of the larvae are greatest when the hybrid larvae are grown on the plant
corresponding to the strain of the mother, rather than the father. For
example, the F-i hybrids where the mother was of the kale-strain grow
larger and pupate sooner on kale than on mustard. Likewise, the Fi
hybrids where the mother was of the mustard-strain grow larger but
not faster on mustard than on kale.
LITERATURE CITED
HOVANITZ, W. aod VINCENT C. S. CHANG. 1962. The effect of vari-
ous food plants on survival and growth rates of Fieru J. Res. Lepid.,
1(1): 21-42.
162
HOVANITZ AND CHANG
/. Res. LepiJ.
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Journal of Research on the ^epidoptera 1(2) ; 163- 168, 1963
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© Copyright
CHANGE OF FOOD PLANT PREFERENCE BY
LARVAE OF PIERIS RAPAE CONTROLLED
BY STRAIN SELECTION, AND THE
INHERITANCE OF THIS TRAIT’
WILLIAM HOVANITZ and VINCENT C. S. CHANG
California Arboretum Foundation, Inc.
Los Angeles State College, Los Angeles
and
University of California, Riverside
It has been shown that larvae which have previously been fed on
a particular food plant are more likely to select that kind of plant than
if they had not previously been fed on it (Hovanitz and Chang, 1962 a
and b). These data have shown that there was a shift in selection of
mustard or of kale according to whether or not the larvae were from a
kale or a mustard strain. Likewise, there was shown a greater selection
for nasturtium by larvae previously fed on nasturtium, and indeed, by
these same larvae, a greater selection for mustard over kale. This change
in selection operates whether the larvae were fed on the changed food-
plant for a long time (several generations) or a short time (part of the
individual larval life) .
In this paper, it is our purpose to test the effect of continued
feeding of the larvae of Pieris rapae for many generations on mustard
and kale by the process of isolating the strains so fed for many genera-
tions, then crossing the parental strains to get Fi individuals and finally
crossing the Fi to get the Fq segregation.
THE STRAINS
The strains of Pieris rapae used in these experiments were derived
originally from two sources, both in the Los Angeles Basin of southern
California. The first, here designated the kale strain, originated with
wild females obtained in a cabbage field (Brassica oleracea var. capitata)
in a truck crop growing area in western Orange County, near Hunting-
ton Beach. The other, here designated as the mustard strain, originated
in the fields of the Los Angeles State and County Arboretum, where in
the spring time Pieris rapae may be found in conjunction with black
mustard (Brassica nigra). These have probably had no recent contact
with cabbage since there are none grown commercially within nine
miles of the area. Before testing in the experiments here described, the
kale strain had passed through more than ten generations in the labora-
1 Aided by a grant from the National Science Foundation, Washington, D. C.
163
164
HOVANITZ AND CHANG
/. Res. Lepid,
tory on kale and the mustard strain had passed through more than six
generations in the laboratory on mustard.
Larvae were selected from each and tested for their seleaion for a
series of plants in the manner indicated in a previous paper (Hovanitz
and Chang, 1962). The plants used in these tests were mustard
(Brassica nigra), kale (Brassica oleracea var. acephala), nasmrtium
(Tropaeolum majus), Isomeris (Isomeris arborea) and Cleome (Cleome
huea).
STRAIN SELECTIONS
The tests were physically carried out in the manner indicated above.
In order to be precise on the nature of any difference to be detected by
these experiments, the tests were conducted in large numbers. Twenty-
five larvae of each strain were used for twenty or forty trials each, giving
a total of six hundred test times for each strain (table I). This was
increased to six hundred sixty for the Fi and nine hundred (with 45
individuals) for the F2.
mustard
kale
nasturtium
Isomeris
Cleome
none
no. of
total
larvae
test
PARENTS
On mustard
364
60.66%
119
19.83%
56
9.33%
18
3.0%
12
2.0%
31
5.16%
25
600
On kale
144
24.00%
354
59.0 %
50
8.33%
21
3.5%
15
2.5%
16
2.66%
25
600
T
M 9x K cf
356
59.93%
141
21.36%
90
13.63%
23
3.48%
39
5.91%
11
1.67%
25
660
(on mustard)
9 d-
(on kale)
328
49.68%
127
19.24%
109
16.51%
33
5.00%
36
5.45%
27
4.09%
25
660
^2
m'9x K d-
491
54.55%
145
16.11%
190
21.4 %
30
3.33%
42
4.66%
2
0.22%
45
900
(on mustard)
K9 X M cf
(on kale)
406
45.11%
146
16.22%
219
24.33%
79
8.77%
39
4.33%
11
1.22%
45
900
Table 1. The comparative mortality and length of larval growth period of
various strains of Pieris rapae.
The differential selection of the plants by larvae of the two selected
strains is clearly made apparent by the curve showing the percentage
selection (fig. 1). The larvae from the mustard bred strain selected
mustard sixty-one percent of the time as compared with kale twenty
percent of the time. The kale strain larvae selected kale fifty-nine
percent of the time as compared with mustard twenty-four percent of
the time. The two strains show an almost complete reversal of their
i(2): i6j-i6S, 1963
PIERIS STRAIN SELECTION
165
preferences with regard to these two plants, their preference being in
the direction of the plant utilized as food for several generations.
The selection by larvae of these two strains for the other plants
concerned was not significantly different. Nasturtium was selected nine
percent of the time by the mustard strain and eight percent of the time
by the kale strain. Isomeris was selected three percent of the time by
each strain, and Cleome about two percent of the time. About five
Fig. 1. Curves showing the percentage selection of various food plants by
larvae of Pieris rapae from a kale and a mustard strain.
percent of the larvae of the mustard strain left test area with no
selection as did about three percent of those of the kale strain. Thus,
there is no difference observable between these two parental strains in
their selection, except with regard to mustard and kale.
THE CROSSES
Adults of the two strains were crossed and the larvae tested for
their preference toward a selection of plants. Since there was a possi-
bility of a maternal influence on the inheritance of the food plant
selectivity hinted at in a previous experiment, reciprocal crosses were
166
HOVANITZ AND CHANG
/. Res. Lepid.
Fig. 2. Curves of the percentage selection of various food plants by larvae of
Pieris rapae comparing the reciprocal F, of the strains shown in figure 1.
made, mustard strain female X kale strain male and vice versa. In each
case, however, the larvae were bred on the plant of the female strain
until tested. It is now known that it would also have been desirable to
have had the reciprocal feeding tests made in addition, as can be seen
below.
The F, larvae obtained, in which the female parent was from the
. mustard strain, showed little difference in selection of plants from the
parent mustard strain except in an increase in selection of nasturtium
and Cleome, neither plant of which was involved in this selection. The
reciprocal Fi larvae obtained from the cross of a kale strain female with
a mustard strain male showed a much greater selection of mustard
than the kale parent strain, and was more like that of either the
mustard parent or of the reciprocal strain indicated (table 1, and fig. 2).
In fact, the results of this cross would indicate that the genes for
mustard selection are nearly completely dominant over those for kale
selection. The cross would also indicate that they are transmitted at
least through the male since a mustard strain male was used with a kale
strain female and the larvae were bred on kale. The fact that the larvae
were bred on kale may be the only reason that the selection was not
higher toward mustard than is indicated. As with the reciprocal cross,
there was a great increase in selection of nasturtium and Cleome
following crossing. This was followed slightly by an increase toward
Isomeris though this may not be significant. There is little doubt from
these data that the selection of food plants is inherited, and that
i(2):i6}-i6S, 196}
PIERIS STRAIN SELECTION
167
"mustard” is dominant over "kale” despite the fact that there is also
superimposed upon this inheritance a selective propensity controlled
by training during the life of the individual, as has been shown before.
The F2 crosses were made in the same manner. F2 larvae of each of
the crosses indicated above were obtained and bred on the food plant
of the mother ( table 1 ) . The tests on these larvae led to basically the
same results as on the Fi larvae with the exception that all percentages
were reduced slightly and the percentages for nasturtium were raised
greatly. Of significance too is the reduction of the larvae which made
no selection at all. The reduction in selection of mustard and kale
appears proportional and related to the increase in the selection of
nasturtium (fig. 3 ) .
DISCUSSION AND CONCLUSIONS
The data indicated in this paper appear to show that selection of
genes for food plant preference occurs over a period of generations
when strains are maintained isolated on particular food plants for many
generations. These genes do not show maternal inheritance. In the
crosses indicated in this paper, genes for selection of mustard over kale
are apparently dominant over the reverse. This is indicated by the fact
that in the Fi of the cross kale strain X mustard strain or the reciprocal,
selection is similar to that shown by the original mustard strain. The
same results are shown for the F^ but they are not so pronounced. Of
great curiosity is the fact that in the Fi there is an increase in selection
for nasturtium, not one of the preferred plants of Pieris rapae ( Flovanitz
and Chang, 1962a). Feeding larvae on nasturtium has shown that there
may be greater selection of this plant developed than would ordinarily
be present. In the present case, however, no selection of this sort is
involved and the increase is significant. This increase is even greater
in the F2 cross than in the Fi, there being a two and one half- to three-
fold increase in selection of nasturtium in the F2 as compared with the
parental strains. A much slighter increase is also indicated for Isomeris
and Cleome, and a decrease for the number of rejects, that is, those that
make no selection. Reasons for this much greater selection for nasturtium
and other plants following hybridization are not known nor can they
even be guessed at intelligently at this time.
SUMMARY
1. Two food plant strains were developed by selection, one on
kale (Bras sic a oleracea) and another on mustard (Brassica nigra).
2. When tested, each of these strains showed a much greater
preference for their accustomed food plant than for any other tested.
Those tested were mustard, kale, nasturtium, Isomeris and Cleome.
3. When these two strains are crossed, the Fi hybrids showed a
preferential selection most like the mustard parent strain. This indicates
that the gene(s) for mustard is dominant over that for kale. There is
168
HOVANITZ AND CHANG
J. Res. Lepid.
Fig. 3. Curves of the percentage selection of various food plants by larvae of
Pieris rapae comparing the reciprocal F2 of the strains shown in
figures land 2.
also indicated a slight increase in selection of the other plants tested.
4. The Fo of this cross showed results similar to the Fi, namely a
selection in favor of mustard rather than kale whether or not the female
parent had originally come from the mustard or the kale strain. Thus,
maternal inheritance is not indicated here even though maternal effects
were indicated on growth rate and mortality data in a previous paper.
5. As had been indicated, feeding the larvae even a short time on
one particular food plant may influence it to have a preferential
selection for that plant.
6. In the F2 there is indicated a strongly increased preferential
selection toward nasturtium which was not selected in any previous
strain. This increase is also indicated slightly toward Isomeris and
Cleome. No reason is advanced at this time for these results.
LITERATURE CITED
HOVANITZ, W. and VINCENT C. S. CHANG. 1962a. The effect of vari-
ous food plants on survival and growth rates of Pieris. J. Res. Lepid.
1(1): 21-42.
1962b. The effect of hybridization of host-plant strains on
growth rate and mortality of Pieris rapae. J. Res. Lepid. 1(2) : this issue.
advertisement
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Volume 1
Number 2
January, 1963
THE J@U^NJAL ©F RESEARCH
©NJ THE LEFIJ©©RTERA\
IN THIS ISSUE
Composition and relative abundance in a temperate zone
butterfly fauna . . . Thomas C. Emmel and John F. Emmel 97
The Argynnis populations of the Sand Creek area, Klamath
Co., Oregon, Part I J- W. Tilden 109
Caterpillar Versus Dinosaur? . . . Theodore H. Eaton, Jr. 114
Geographical distribution and variation of the Genus Argynnis
I. Introduction
II. Argynnis idalia William Hovanitz 117
The relation of Pieris virginiensis Edw. to Pieris napi L.
Species formation in Pieris? Wiliam Hovanitz 124
The male genitalia of some Co lias species . Bjorn. Petersen 135
The effect of hybridization of host -plant strains on growth rate
and mortality of Pieris rapae
William Hovanitz and Vincent C. S. Chang 157
Notice-editorial 162
Change of food plant preference by larvae of Pieris rapae
controlled by strain selection, and the inheritance of this trait
William Hovanitz and Vincent C. S. Chang 163
@F
OIMJ THE LEFI1P@PTER1A
MeSfeii
Bi2!!*S*****»»*«'*" .
...
^^■rs****** ***•*•»
!psr.i:j2:r*fe;
us:sss:2s;*.2ui,.
K«sjssK2.»:2i;ir
Volume 1
Number 3
March, 1963
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Journal of Research on the Lepidoptera 1 ( 3 ) :169-182, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 196}
SELECTION OF ALLYL ISOTHIOCYANATE BY
LARVAE OF PIERIS RAPAE AND THE
INHERITANCE OF THIS TRAIT'
WILLLIAM HOVANITZ and VINCENT C. S. CHANG
California Arboretum Foundation, Inc., Arcadia,
Los Angeles State College and
University of California, Riverside
Larvae which have previously been fed on a particular food
plant are more likely to select that plant than if they had not previously
been fed on it (Hovanitz and Chang, \9G2a and ^). It was shown that
larvae which were fed on kale are more likely to select kale, and larvae
which were fed on mustard are more likely to select mustard. Also,
larvae which have previously been fed nasturtium have a higher
selection on that plant than if they had not been fed on it. This
change in food plant selection operates both if the adaptation comes
over a period of several generations, or if only for a short time in the
individual larval life. Later, it was shown that strains selected for
several generations on each of two plants, kale and mustard, had strong
selection for their respective food plants ( Hovanitz and Chang,
1963^^ and b). Individuals from these two strains were then crossed to
get the Fi generation, and these were crossed to get the Fo generation.
The inheritance of this trait appears to be multi factorial but because
of the nature of the testing, this point may be open to question.
The data to be presented in this paper were obtained in much the
same manner as in the previously mentioned papers (Hovanitz and
Chang, 1963<? and b) with the exception that instead of testing the
selection of the larvae for the plants themselves, they were tested for
their selection of various concentrations of black mustard oil (allyl
isothiocyaoate ) . Tests were made on the parental larvae as well as the
Ft and the F2.
THE STRAINS
The strains of Pieris rapae used in these experiments were derived
from two original sources. The first, the kale strain, originated with wild
females obtained in a cabbage field {Brassica oleracea var. capitata)
in a truck crop growing area near Huntington Beach in Orange County,
California. The other, the mustard strain, originated in the fields of the
'Aided by a grant from the National Science Foundation.
169 SlllTflSIJNlW
mSTlTUTiaH
APR1819®
170
HOVANITZ AND CHANG
/. Res, Lepid.
Los Angeles State and County Arboretum where P. rapae is found on
black mustard (Brassica nigra). Before testing in the experiments here
described, the kale strain had passed through more than ten generations
in the laboratory on kale {Brassica oleracea var. acephala) and the
mustard strain had passed through more than six generations in the
laboratory on black mustard.
EXPERIMENTAL SET-UP
The tests of the larvae were made in much the same manner as has
previously been described (Hovanitz and Chang, 1962b) except that
in place of the plants to be selected, at various points surrounding the
flat, filter papers wetted with various solutions of allyl-isothiocyanate
were placed. Preliminary tests had indicated that the larvae were
attracted to dilutions of the mustard oil at 10"^ to 10"'^ so the tests were
made at these dilutions. Metal mosquito netting was cut in a circular
arrangement to serve as a base for the filter paper. These discs of metal
netting were placed on top of glass flasks partially buried in soil, secured
to corks placed in the flasks and arranged in various parts of the flat
according to the disposition shown in Figure 1. The filter paper was
then set up top of the metal netting and wetted with distilled water
solutions as shown. On one of these, distilled water only was placed so
as to serve as a control for the others. To prevent the accumulation of
mustard oil concentrations, the solutions were alternated around the
flat. The set-up of the experiments is shown diagrammatically in Figure 1
and photographically in Figure 2. The larvae were set in the center of
the flat and allowed to select any of the six test papers or to leave the
flat without selection. It should he noted that the test papers were above
the level of the larvae and therefore that the heavier-than-air mustard
oils would drop down to their level. After each trial, the soil in the flat
was mixed to prevent the origin of "trails.” It should also be noted that
it would be much easier for the larvae to leave the flat without any
selection except for one factor, namely, that these larvae have a tendency
to climb upwards when at all possible. Experience with lepidopterous
larvae has shown that descent is usually made by means of dropping
down on a silken thread, rather than by walking while ascent is made
by walking. This accounts for the selection of the "water” alone just
as frequently as they leave the flat without any selection at all.
The mustard oil used in these experiments was a natural product
obtained from black mustard seed by compression and distillation. It
is available commercially from the Fritsche Brothers firm. Allyl iso-
thiocyanate is the principal component of the oil obtained from Brassica
nigra; other mustard oils are found in other plants and all are esters of
isothyocyanic acid. Brassica nigra seeds contain a glucoside, potassium
myronate or sinigrin, which undergoes fermentation change due to the
enzyme myrosin present in the seeds.
(})ti69-ii2, 196}
ALLYL ISOTHYOCYANATE
171
Fig. L The set-up for the larval tests. The larvae are placed in the
center of the flat from where they are enabled to go in any direction toward
one of the dilutions of mustard oil, or to leave the flat. The circles show the
disposition of the filter papers which have upon them dilutions of mustard
oil as indicated.
THE TEST SELECTIONS
Test selections were made by using a number of larvae for several
times to get a combined sample. The number of individuals used
depended in part on those available, but ranged from fifteen to forty-five
giving a total number of tests from three hundred to nine hundred.
These data are given in Table 1.
The parent strains were tested first. The mustard strain was tested
with twenty -five larvae 20 or 40 times each giving a total of seven
hundred trials. Of these, only nineteen or 2.7 percent made no selection
and left the flat. Only twenty-four or 3.4 percent selected the filter
paper with distilled water only on it. The others all selected the mustard
oils, from a low of nearly 10 percent for the 10"’ to a high of nearly
thirty percent for the 10"^. These data are shown diagrammatically in
Figure 3.
172
HOVANITZ AND CHANG
/. Kes. Lepid.
Fig. 2: Photograph showing the surface of the experimental flat, the
flasks with filter paper upon them and a larva raising its foreparts. The rules
are not present during an experiment and are indicated only for the purpose
of illustrating the dimensions between the soil surface, the filter paper and
the larvae.
for different concentrations of allyl isothiothiocyanate
(black mustard oil).
1(3)^169-182, 1963
ALLYL ISOTHIOCYANATE
173
5
174
HOVANITZ AND CHANG
/. Res. Lepid,
The kale strain was also tested with twenty-five larvae for a total
of five hundred eighty trials. The highest selection of these larvae was
toward the concentration of 10"®, larvae going to this concentration in
thirty-four percent of the trials. This result is significantly different
from the result for the kale larvae indicated above, in which the highest
selection was toward the 10"® concentration. These data can be com-
pared diagrammatically in Figure 3. There is little doubt that the larvae
of the kale-bred stock prefer the mustard oil at a lower concentration
than do those of the mustard strain. It is probably not coincidental that
the taste of the leaves of mustard is stronger than is that of kale (or
cabbage) since the significant taste of these plants is due to allyl or
other isothiocyanates. These data would seem to indicate ( 1 ) that the
concentration of taste or smell-detectable mustard oils in the leaves
of mustard is higher than that in kale, (2) that the larvae feeding on
mustard have become adapted in some way to a higher concentration
of these oils, and (3) that due to this adaptation, the "mustard” larvae
have a greater selection of the higher concentration of mustard oil than
the lower. It has been shown in another paper (Hovanitz and Chang,
1963b) that these two strains also select their own plants over the
others when given a free choice. This preferred selection may be due
wholly or in part to the difference in available allyl or other isothiocya-
nate in their leaves.
THE CROSSES
The crosses between the two food plant strains were carried out by
making reciprocal hybrids. In the one case, the kale strain was used as
the female parent and in the other case, the mustard strain. The hybrid
larvae resulting from these crosses were fed on the food plant of the
mother, since the mother was allowed to lay on its own preferred plant.
The selections made by the hybrid larvae in both crosses favored
the concentration 10"^, exactly half way between the selections made
by the respective parent strains ( Fig. 4 ) . The frequency of selections
for the kale female Fi toward the 10"^ concentration was 34 percent as
compared with 30 percent for the mustard female Fi.
Previous data have indicated that the immediate larval food plant
eaten by the larvae had an effect on the food plant selections made by
these larvae. If this held true for the selections of the mustard oils also,
then the larvae of the kale female Fi, having been fed on kale during
their life, and the larvae of the mustard female Fi, having been fed
on mustard during their life, should have a preference for their respec-
tive plant. Data from Table 1 (Hovanitz and Chang, 1963b) indicate
that this is so. Mustard female Fi larvae selected mustard nearly 60
percent of the total selections as compared with nearly 50 percent selec-
tions for the kale female Fi larval selections. It should follow, if our
PERCENT SELECTION PERCENT SELECTION
i(3):i69-az, 196)
ALLYL ISOTHIOCYANATE
175
Fig. 3: Histograms illustrating the selections made by the larvae from
the parental strains, comparing the kale and the mustard strains.
PERCENT SELECTION PERCENT SELECTION
176
HOVANITZ AND CHANG
/. Kes. Lepid.
Fig. 4: Histograms illustrating the selections made by the Fi
from the reciprocal crosses between the kale and mustard strains.
larvae
PEHCEMT SELECTiON PERCENT SELECTION
(3);i6^-ii2, 1^63
ALLYL ISOTHIOCYANATE
177
Fig. 5. Histograms illustrating the selections made by the Fa
from the reciprocal crosses between the kale and the mustard strains.
larvae
178
HOVANITZ AND CHANG
/. Res. Lcpid.
assumptions are correct concerning the relationship between mustard
oil concentration and food plant selectivity, that the mustard female Fi
larvae should select mustard oil concentrations at a higher level than
the kale female F] larvae. That this is so can be seen by study of the
histograms (Fig. 4) and the following calculations. Selections made
by the mustard female Fi larvae which are less than 10"^ (that is
10"^ or ‘^) are 31.4 percent of the total as compared with 24.6 percent
of the total for the kale female F, larvae. This shows that the larvae of
the mustard female Fi select a higher concentration of mustard oil than
those of the kale female Fi even though the modal class in both cases
is at 10"^. This difference is most probably attributed to the fact that
the immediate F, larvae had been fed on the food plant of the mother,
rather than that there was any genetic effect of maternal inheritance.
Two sets of Fq have been bred, corresponding to the two sets of Fi.
These two sets of Fi are the reciprocal crosses mentioned above. The Fq
from the mustard female Fi was continued on mustard for the food
plant of the F2 larvae, and the F2 from the kale female Fi was con-
tinued on kale. It would be expected, therefore, that each of these
strains might diverge slightly in their selection according to the pattern
of the larval foods. This result did not materialize.
The data indicate that the Fq of both strains still have a selective
propensity favoring the dilution, 10"^, thus having the same modal
class as the F 1 ( Fig. 5 ) . This is the expected genetic result for multiple
factor inheritance. The second result expected for multiple factor in-
heritance is a wider range of variation in the F2 than the Fi. This
result is not shown by a wider range per se since no wider range of
dilutions were used, but rather it is shown by a greater dispersion of
the selections inside the range tested. This would give a greater standard
deviation for the F2 than for the corresponding Fi. The greater disper-
sion is very clear by study of the histograms, rearranged so that the
mustard strains are on one page and the kale strains are on another
(Figs. 6 and 7). The differences between the Fi and the F2 are striking.
These results indicate clearly that the factors for food plant selection in
Pieris rapae which have been separated in the two strains here tested
are genetic and behave in a way comparable with polygenic inheritanp.
It also indicates that the populations of Pieris rapae existing in the wild
are differentiated into genetically distinct populations, differing by genes
which control the selection of food plants. These differences could
account for a mechanism needed for species isolation (Hovanitz 1963c).
DISCUSSION AND CONCLUSIONS
The data indicated in this paper show that the strains which have
been selected for particular food plant preferences over a period of
several generations, and which when tested prefer the food plant upon
which they have fed, also differ in their selection (preference) for vari-
ALLYL ISOTHIOCYANATE
179
ous concentrations of mustard oil (allyl isothiocyanate). That these
differences are genetic is apparent not only from the preceding data
on food plant preferences but also on the data on mustard oil preferences
introduced in this paper. Crossing of the two strains discussed here gives
an F, which has a selection exactly intermediate between the two par-
ental strains. The results are similar whether the kale strain contributes
the maternal parent or whether the mustard strain contributes the
maternal parent. There is, however, a greater selection of the higher
concentrations in the strain fed on mustard than the one fed on kale.
The F2 from these strains is similar to the Fi except for the wider
dispersion of the selections.
In the testing of these strains for food plant preferences, it was
found that the larvae developed a preference for nasturtium in the Fi
and F2. No reason can be deduced for this strange event based upon the
present results unless it be that nasturtium emits at a concentration of
mustard oil of the same attractive power as 10"^ allyl isothiocyanate or
some other mustard oil of attractive significance. The odor of nasturtium
does not seem to the human to be the same as that of allyl iosthiocyanate.
However, it is possible that to the human, the odor of some other sub-
stance masks the odor of the attractive mustard oil, but that this odor
does not do so in the larvae which are able to detect it.
Thorsteinsen (1953) has made the statement that larvae of the
diamond-back moth, Plutella macuUpennis (Cut.) are attracted to
the odor of ally! isothiocyanate from a distance but are repelled by it
when closeup so that they will not eat substances having this oil. Instead,
they will eat substances with the corresponding glucoside, potassium
myronate, but are not attracted to this substance from a distance. Since
the glucoside is converted into the allyl isothiocyanate in the plant by
a process of fermentation, it is probable that the attraction is really for
the isothiocyanate not the glucoside, but that the isothiocyanate is pro-
duced slowly and in small quantities by the process of enzymatic
fermentation. Indeed, Thorsteinson even indicates "in some experi-
ments the addition of allyl mustard oils slightly increased feeding on
media containing sinigrin.” Wolfrom (I960) points out that "while
the exact biological function of the plant glycosides is not established,
it is probable that their formation provides the plant with a means of
storing, in a harmless form, toxic and physiologically active materials
which may be liberated by enzymes, in small quantities, when required.”
It seems not only possible but highly likely that the attraction differ-
ences noted by Thorsteinson are based upon the low concentration of
allyl isothiocyanate rather than upon the presence of the glucoside
sinigrin directly. Thorsteinson in fact admits this possibility in his com-
ment, "On the other hand, it is possible that infinitesimal amounts of
mustard oil vapor emanating from the leaves [by enzymatic action of
myrosin on sinigrin] may stimulate the olfactory sense which is char-
acteristically extremely sensitive in insects.” Section in brackets not
PERCENT SELECTION PERCENT SELECTION
180
HOVANITZ AND CHANG
/. Res. Lepid.
Fig 6. Histograms illustrating the selections made by the Fi and F:;
larvae, comparing these larvae of the kale female crosses.
PERCENT SELECTION PERCENT SELECTION
(}):i69-i82, 196}
ALLYL ISOTHIOCYANATE
181
Fig. 7. Histograms illustrating the selections made by the Fi and F2
larvae, comparing these larvae of the mustard female crosses.
182
HOVANITZ AND CHANG
/. Res. Lepid.
Thorsteinson’s. Further testing of feeding responses of Plutella or
Pieris larvae at a low mustard oil level are required to be certain of
this differentiation. Presently published data of the present, or of other,
authors are definitely not conclusive. Tests of this sort may be very
difficult to make due to the high volatility of the mustard oil. There is
no doubt that the use of the glucoside, rather than the oil, makes easier
the maintenance of a low level of the oil vapor in the vicinity of the
food. Our own data comparing the egg laying of the adults which are
to be published later shows that the glucoside and mustard oil released
from crushing seeds is more attractive than the mustard oil alone.
Verschaffelt (1910) was the first to study the relationship between
the food plant choice of insects, plant odor and the chemistry of the
relationship. He supposed that the larvae were attracted to the mustard
oils present in the food plants but did not know whether the larvae
differentiated between different oils or not. Dethier (1941) showed
the selective effect of various odiferous oils toward lepidopterous larvae.
Johansson (1951) showed that Pieris hrassicae larvae preferred food
plants to which they had already become accustomed by previous
eating. This result has also been shown by us (Hovanitz and Chang
1962b and 1963a) on Pieris rapae.
LITERATURE CITED
DETHIER, V. G. 1941. Chemical factors determining choice of food pianis
by Papilio larvae. Amer. Nat. 75: 61-75.
HOVANITZ, WILLIAM and VINCENT C. S. CHANG. 1962a. The effect
of various food plants on survival and growth rate of Pieris. /, Res.
Lepid. 1.(1): 21-42.
— — ■. 1962b. Three factors affecting larval choice of food plant.
J. Res. Lepid. 1(1): 51-62^
— — — ■. 1963a. The effect of hybridization of host-plant strains on
growth rate and mortality of Pieris rapae. J. Res. Lepid. 1 (2) : 157-161.
• — . 1963b. Change of food plant preference by larvae of Pieris
rapae controlled by strain selection, and the inheritance of this trait.
/. Res. Lepid. 1 (2) ; 163-168.
. 1963c. The relation of Pieris virginiensis Edw. to
Pieris napi L. Species formation in Pieris? /. Res. Lepid. 1 (2) ; 124-134.
JOHANSSON, ARNE SEMB. 1951 The food plant preference of the larvae
of Pieris hrassicae L. Norsk Ent. Tid. B. VIII, 2.4-5: 187-195.
THORSTEINSON, A. J. 1953. The chemotactic responses that determine
host specificity in an oligophagous insect {Plutella maculipennis (Curt.)
Lepidoptera ) . Canad. J. Zool. 31: 52-72.
VERSCHAFFELT, E. 1910, The cause determining the selection of food in
some herbivorous insects. Proc. Acad. Sci. Amsterdam 13(1): 536-542.
WOLFROM, M. L. I960. Natural glycosides. Encyclopaedia Britannica
10: 448..
Journal of Research on the Lepidoptera 1 (3 ); 183=190, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 1^6}
BIOLOGY OF THE CEANOTHUS STEM-GALL MOTH,
PERIPLOCA CEANOTHIELLA (Cosens)
With Consideration of its Control*
J. ALEX MUNRO
Los Angeles State and County Arboretum
Arcadia, California
The stem-gall moth, Periploca ceanothiella {Cosens), has
received but slight attention considering its importance as a pest of
ornamentals on the genus Ceanothus. The genus Ceanothus, according
to Van Rensselaer and McMinn (1942), comprises some 55 species
native only to North America. Of these, some 44 species, and a number
of horticultural varieties or selections, are grown in the Pacific Coast
states. Many of them exhibit rather colorful bloom, ranging from the
white flowers of C. americanus (New Jersey Tea) to the deep blue,
lilac -like clusters of C. thyrsifloms (Blue Blossom) and the pink of
some selections. They occur mainly as shrubs and are used in land-
scape plantings and roadside beautification. Some, such as C. gloriosus
exaltatus (Point Reyes Creeper) and C. griseus horizontalis (Carmel
Creeper) are used as attractive groundcovers because of their low
growing habit.
The distribution of the gall moth probably corresponds to the range
of its host. Hodges (1962) in reviewing the genus Periploca mentions
having examined specimens of P. ceanothiella collected in California,
Kansas, New York, Ontario (Canada), and Texas. His review shifted
this insect from the genus Stagmatophora, which it formerly occupied,
to its present position in the genus Periploca.
DEVELOPMENT
This insect normally overwinters within its gall in some stage of
larval development. Reference to Table 1 shows pupation to be barely
under way in April and to continue, with moth emergence, throughout
spring and early summer. Emergence, however, may begin as early as
January from mature larvae brought in from outdoors and maintained
under room temperatures.
The small, dark-colored moths (Fig. 1) begin egg laying within a
few days following emergence. They deposit their small (0.5 mm) white
■^Presented at the annual meeting of the Entomological Society of America, Phoenix, Arizona —
December 3-6, 1962.
183
184
MUNRO
/. Res. Lcpid.
TABLE 1. Status of overwintered Periploca ceanothiella galls.
PLACE 6 DATE
PERCENT GALLS
1962
Number of
galls exam
With
Larvae
With
Pupae
With
Parasites
Emerged
Arcadia
April 9
88
87
2
11
0
May 18
64
60
21
15
4
June 7
142
23
44
19
14
July 26
104
2
6
3
89
Claremont
April 25
85
52
28
18
2
May 16
80
28
22
11
39
June 14
76
19
26
15
40
La Canada
April 17
66
94
0
6
0
May 22
70
62
4
19
15
Santa Barbara
April 14
141
95
1
4
0
eggs on the underside of leaves, in leaf axils, and along the twigs. The
eggs required 10 days for hatching under indoor temperatures of about
70° F. The newly hatched larvae move directly into the terminal growth
where they penetrate the buds and inflorescences to cause the swellings
or galls. Here the larva remains, feeding and growing on the inner
tissue, until reaching full growth of 5 to 6.5 mm by the following
spring or summer.
Before pupation begins, the larva cuts an exit hole partially through
the side of the gall to leave only a thin film of bark which covers the
entrance until time for the moth to emerge. This "window” or blocked
entrance provides protection for the occupant against some enemies
until it is ready to escape from the gall cavity. The pupation stage is
spent within the gall and requires 24 days under room temperatures of
about 70 °F. Although the gall moths may emerge over a prolonged
period of spring and summer, only one brood or annual generation was
observed.
The pest has a stunting effect on the new growth. The galls may
vary greatly in abundance, ranging from only an occasional gall to 20
or more per lineal foot of branch and twigs in heavy infestations. Some
(^)-.iS3-i90, 19^3
GALL MOTH
185
of the twigs may be killed outright from the injury. The most serious
damage, however, was to the bloom. (Fig. 2). It was not unusual to
see the flower clusters reduced to about 25% of their normal size,
particularly where the gall larvae were present in the inflorescenses.
In no instance was more than one larva found per gall. In appear-
ance, the gall is somewhat spindle-shaped and averaged about 13mm
in length by 6mm diameter. (Fig. 3) This is about three times the
diameter of the twig at point of junction with the gall.
PARASITES
The role of three naturally occurring ichneumonids in checking
the stem gall is indicated in Table 1. All three, Fristomems haumhoferi
Cush., Scambus aplopappi (Ashm.), and Apistephialtes nucicola
(Cush.), were observed mainly during the spring months. No hyper-
parasites were observed. Identifications were made by L. M. Walkley
of the U.S. National Museum Staff. Muesebeck, Krombein, and Townes
(1951) mention these ichneumonids as parasites of certain micro-
lepidopterous larvae that tunnel in stems. Of the three parasites, P.
haumhoferi was the most commonly encountered, and was the only
one previously recorded as a parasite of this stem gall insect. The toll
taken by the parasites ranged up to 19% of the gall-larvae. Only the
mature larvae were attacked by the parasites.
NON-SUSCEPTIBILIl’Y IN CEANOTHUS
Examination of Ceanothus collections at the Los Angeles State
and County Arboretum in Arcadia, Rancho Santa Ana Botanic Garden
at Claremont, and the Santa Barbara Botanic Garden at Santa Barbara
(all southern California locations) showed varying degrees of non-
susceptibility to the stem galls. It was an ideal way to check on this
factor inasmuch as the Ceanothus species and varieties growing in each
collection were equally exposed to the pest. The results presented in
Table 2 would suggest the use of non-susceptible Ceanothus in plant-
ings wherever practicable.
CHEMICAL CONTROL
Inasmuch as spraying of the Ceanothus with DDT had given little
or no apparent control of this pest it was decided to test the systemic
insecticide, dimethoate, for the purpose. A commercial formulation
(Cygon) containing 43.5% dimethoate was mixed at the rate of one
pint of this concentrate to 100 gallons of water and applied as a foliar
spray to Ceanothus plantings in Arcadia on June 27, 1962. Examina-
tion made after one week following application showed that 92% of
186
MUNRO
/. Res. Li pid.
TABLE 2, Occurrence of Periploca ceanothlella on various species, varieties
and horticultural selections of Ceanothus.
HEAVY OCCURRENCE;
Ceanothus griseus
C, griseus horizontalis
MODERATE OCCURRENCE;
C. cyaneus
C, thyrsiflorus
C, 'Ray Hartman*
C, 'Marie Simon'
LIGHT OCCURRENCE:
C. arboreus
C, oliganthus
C, divers if olius
C. lemmonii
C, integerrimus
C, leucodermis
C, lobbianus
C, 'Treasure Island*
C. 'Sierra Blue*
C, 'Royal Blue*
C, 'Mountain Haze*
C, 'Mary Lake'
C. 'Concha*
NO OCCURRENCES
C, americanus
C, parry i
C. papillosus
C, impressus
C, foliosus
C. insularis
C. cuneatus
C« ramulosus fascicularis
C, gloriosus
C. gloriosus exaltatus
C. rigidus albus
C. purpureus
C. prostratus
C. verrucosus
C. spinosus
C, jepsonii
C, raasonii
C« megacarpus
C, greggi perplexus
C, 'Blue Cloud*
C. 'Lester Rowntree*
the gall larvae were killed. The spray, however, had no appa'rent effect
on the mature gall larvae (which had ceased feeding), the pupae, or
the parasites, P. baumhoferi and A. nucicola; adults of these normally
emerged from galls collected from the treated areas. Effect of the treat-
ment on 5’. aplopappi was not observed. The parasites developed only
in the mature gaU larvae, and therefore it is unlikely that they had had
any contact with the insecticide. In addition to checking the stem gall
infestation, the dimethoate also controlled the Ceanothus leaf miner,
Nepticula ceanothi Braun, thrips, and an infestation of mites which
were troublesome at the time. The spray used at this rate had no ob-
servable phytotoxic effect on the Ceanothus.
A similar application made on September 14, 1962 showed only
74% mortality of the gall larvae. This lower kill may have been
due to the greater proportion of older, possibly more resistant, larvae at
this time than at the June 27th spraying.
A soil drench of the dimethoate, made at six times the strength of
the foliar spray, applied around the base of Ceanothus shrubs in Pasa-
dena on July 11, 1962, resulted in 70% mortality of the gall larvae, but
196}
GALL MOTH
187
M E T R I C
.j_,.
J_j_j.
1 1, i
S Y S T E .M
Fig. 1. The adult moth, Periploca ceanothiella (Cosens), at bottom
right; larva and pupa at left; larva in cut gall at top.
188
MUNRO
/. Res. Lepid.
I>
i(}):iS}-i90, 196}
GALL MOTH
189
Fig. 3. Stem galls from which the moths have emerged; note the exit
hole in the encircled gall. The twig at the right has been completely killed by
the larvae. Note that the galls occur at the beginning of each period of stem
growth, and that in the twig at the left, two different years’ galls are shown.
190
MUNRO
/. Kcs. Lcpid.
caused burning and subsequent defoliation wherever the drench came
in contact with the low-growing foliage. Examination of galls from
untreated Ceanothus coinciding with observation on the treated shrubs,
showed natural mortality of less than one percent, and of no significance.
Limited observations on Ceanothus growing in the wild showed the
stem-galls to be less abundant than on Ceanothus grown under culti-
vation. This might indicate that irrigation and other care which these
ornamentals received under cultivation makes them more attractive
to this gall insect.
SUMMARY
The stem-gall moth, Periploca ceanothiella ( Cosens ) , is a pest of
ornamental shrubs in the genus Ceanothus. The moths emerge from the
galls and lay their eggs throughout spring and early summer. There is
but one generation or annual brood. Three naturally occurring ichneu-
monid parasites accounted for up to 19% mortality of the maturing
gall larvae. No hyperparasites were observed. Examination of 40 species
and varieties of Ceanothus growing in mixed collections at several
southern California locations showed 2 with heavy occurrence of the
galls, 4 with moderate occurrence, 13 with light occurrence, and 21
with no occurrence. This would suggest the planting of the more
non-susceptible varieties to alleviate or avoid the problem where
practicable. Dimethoate used as a foliar spray proved highly effective
against the immature gall larvae, as well as the leaf miner, thrips, and
mites, but had no apparent effect on the mature gall larvae and pupae,
or the parasites. Limited observations indicated the galls to be more
troublesome on Ceanothus growing under cultivation than in the wild.
REFERENCES CITED
VAN RENSSELAR, M., and H. E. McMINN. 1942. Ceanothus. Gillick
Press, Berkeley, California. 308 pp.
MUESEBECK, C. F. W., K. V. KROMBEIN, and H. K. TOWNES. 1951.
Hymenoptera North of Mexico Synoptic catalog. Monograph No. 2, U. S.
Department of Agriculture, Washington, D.C. pp. 186, 188 and 387.
HODGES, R. W. 1962. A review of the genus Periploca with descriptions
of nine new species. Pan-Pacific Entomologist. 38: 83-89.
Journal of Research on the Lepidoptera
1(3):19M93, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 15)63
LARVAL FOOD-PLANT RECORDS FOR
SIX WESTERN PAPILIOS
JOHN F. EMM£L and THOMAS C. EMMEL
Stanford University, California and Reed College, Portland, Oregon
Reported on here are some of our recent observations on
natural and laboratory foodplants used by Papilio eurym-edon Lucas,
P. indra indra Reakirt, P. indra pergamus Edwards, P. bairdii Edwards,
P. oregonius Edwards, and P. rudkini Comstock. Some previously re-
ported foodplants whose authenticity has been doubted are shown to be
true larval foodplants, while several new plant records for these species
are reported for northern Oregon and California populations.
1. Papilio eurymedon.
After a thorough review of available records, Brower (1958; 1962,
in litt.) concluded that Pmnus is not a foodplant of this species. The
present authors (1962), however, recorded this plant as a possible
foodplant for P. eurymedon, and several observations of eurymedon
using Prunus seem to be worthy of publishing in view of the previous
conflicting evidence.
While collecting in the vicinity of Frazier Park, Kern County,
California, on June 30, 1962, the senior author observed what appeared
to be a female Papilio eurymedon fluttering over a small bush of Prunus
ilicifolia. This observation led to a search of the plant, which yielded one
second-instar Papilio larva. This larva was brought back to the labora-
tory and was successfully reared on Prunus lyoni. It pupated on August
6 and on August 22, it produced a male eurymedon.
Noel McFarland (1962, in litt.) has also found Prunus ilicifolia,
as well as Rhamnus crocea, to be foodplants of Papilio eurymedon. At
Oak Pass in the Santa Monica Mountains of Southern California, he
states: "Prunus ilicifolia ... is the only plant I have ever found them
on (beyond second instar) in the wild. I have often collected eggs and
first instar larvae on Rhamnus croceaP
At least in California, then, Prunus ilicifolia seems to be a natural
and fully satisfactory foodplant for Papilio eurymedon.
2. Papilio indra indra.
In his Butterflies of North America (1897), Edwards stated that
Artemisia dracunculoides (Compositae) was a foodplant of P. indra in
Colorado. Kent Wilson (1961) apparently used this record in the most
recent publication of foodplants for this species. However, in 1918 J. C,
Hopfinger- reported that he had never found indra larvae on A. dracun-
culoides. He did find black Papilio larvae (very probably indra) on an
191
192
EMMEL AND EMMEL
/. Res. Lepid.
"umbelliferous species,” on which he also found larvae of P. zelicaon.
These black "indr a” larvae would not accept A. dracunculoides when
transferred to it.
. As we reported in 1962, the foodplant of P. indr a in the Sierra
Nevada is Pteryxia terebinthina ( formerly Cymopterus terehinthinus ) ,
There are many botanical records of this plant (Dr. Mildred Mathias,
personal communication) for the area around Brewster, Washington, and
the "umbelliferous” foodplant found by Hopfinger may well have been
this species. Don Eff (1962, in litt.) reports the foodplant of indr a in
the Front Range of Colorado to be Harbouria trachpleura (Umbelli-
ferae) .
3. Papilio indr a pergamus.
The first known foodplant of this subspecies of indra was found by
Comstock (1928); this was Tauschia parishii (Urn belli ferae) in the
San Gabriel Mountains.
Fred Thorne (1962, in litt.) has found pergamus eggs and larvae
on Tauschia arguta and Lomatium lucidum (Umbelliferae) on Tecate
Peak, San Diego County, California.
4. Papilio bairdii.
Edwards (1893, 1898) found that carrot (Daucus carota) was
somewhat acceptable to bairdii larvae, while the larvae "thrived” on
fennel {Foeniculum vulgar e) . However, Brown (1957) states that
these two plants are unacceptable to bairdii.
On July 27, 1962, the senior author collected 6 fifth-instar larvae
and 2 fourth-instar larvae of P. bairdii on Artemisia dracunculoides
(Compositae) at Barton Flats, San Bernardino County, California. In
the laboratory, these larvae immediately accepted fennel when placed
on this plant. Fennel and this Artemisia were eaten with no preference
for either plant. These larvae pupated, and a male and female adult
pair emerged on September 9, 1962.
5. Papilio oregonius.
On September 1, 1962, both authors collected larvae of P. oregonius
at Heppner Junction (Gilliam County) along the Columbia River,
Oregon. These larvae ( 1 second-instar, 2 fourth-instar, and 3 fifth
instar larvae) were found on Artemisia dracunculoides. In the labora-
tory, they fed readily on fennel. It was of possible biochemical interest
to note that the odors of crushed leaves of these plants were similar.
6. Papilio rudkini.
The natural foodplant of this species is Thamnosma montana
(Rutaceae). But this Papilio has also been found on Daucus carota
(Umbelliferae) in Yuma, Arizona (Bauer, 1955).
In April of 1962, the senior author collected ten larvae of Papilio
rudkini on Thamnosma montana in Sentenac Canyon, San Diego
County, California. These larvae were transferred to fennel in the
laboratory, which was fairly acceptable to them (20% mortality). All
refused to eat Citrus, which is very acceptable to Papilio zelicaon.
PAPILIO FOOD PLANTS
193
SUMMARY OF NATURAL AND LABORATORY FOODPLANTS
RECORDED IN THIS PAPER
N— natural foodplant; L=:acceptable as laboratory foodplant.
Papilio eurymedon Rosaceae
Pmnus ilicifolia (N)
Prunus lyo-ni (L)
Rhamnaceae
Rhamnus crocea (N)
Papilio indra indra Umbelliferae
Pteryxia terehinthina (N)
Harbouria trachpleura (N)
Previously-recorded Artemisia dracnticnloidcs is dropped in this paper from the group of known
foodplants of P. indra indra.
Papilio indra pergamus
Papilio bairdii
Papilio oregonius
Papilio rudkini
Umbelliferae
Tauschia parishii (N)
Tauschia arguta (N)
Lomatium lucidum (N)
Compos itae
Artemisia dracunculoides (N)
Umbelliferae
Daucus car Ota (L)
Foeniculum vulgare (L)
Compositae
Artemisia dramunculoides (N)
Umbelliferae
Foeniculum vulgare (L)
Rutaceae
Thamnosma montana (N)
Umbelliferae
Daucus carota (N, buti
introduced plant)
Foeniculum vulgare (L)
LITERATURE CITED
BAUER, DAVID L., 1955. Notes on the Papilio machaon complex in Arizona.
Lepid. News 9: 7-10.
BROWER, LINCOLN P., 1958. Larval foodplant specificity in butterflies
of the Papilio glaucus group. Lepid. News 12: 103-114.
BROWN, F. MARTIN, 1957. Colorado Butterflies, p. 215.
COMSTOCK, JOHN ADAMS, 1928. Records of the Lorquin Entomological
Society. Bull. Southern Cal. Acad. Sci. 27: 73-
EDWARDS, WILLIAM H., 1893. Notes on a polymorphic Papilio. Canad.
Ent. 25: 253-254.,
, 1897j Butterflies of North America, 3 vols.
- — j 1898. Further observations on Papilio bairdii Edwards.
Canad. Ent. 30: 11.
EMMEL, THOMAS C., & JOHN F. EMMEL, 1962. Ecological studies of
Rhopalocera in a High Sierran community- — Donner Pass, California.
L Butterfly associations and distributional factors. Journ. Lepid. Soc.
16: 23-44.
HOPFINGER, J. C., 1918. Notes on Papilio indra Reakirt. Ent. News 29:
354-5.
WILSON, KENT H., 1961. Family Papilionidae. pp. 30-50. In Ehrlich,
Paul R. and Anne H., 1961. How to Know the Butterflies, Dubuque,
Iowa: Brown, 262 pp.
1 (3) :194, 1963
Journal of Research on the Lepidoptera
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright
COLIAS PHILODICE IN CHIAPAS, MEXICO
THOMAS C. EMMEL
Reed College, Portland, Oregon
COLiAs PHILODICE GoDART has the widest continuous range of any
Goliad in North America. The species is found from the northern Arctic
southward to the edge of desert areas in the Owens Valley of eastern
California, the highlands of northern Arizona, and the Cochise and Rio
Grande River valleys of New Mexico and Texas. In addition, three
isolated populations of the species are known in the central mountains
of Guatemala (Hovanitz, 1950a). The present note extends the known
distribution of the species to include the republic of Mexico, and gives
the frequency of white dimorphic females in the single population
found.
The Mexican population of C. philodice was discovered in a long
valley about 12.5 kilometers southeast (on Highway 190) from the
plaza in San Cristobal de las Casas, central Chiapas. The valley is about
a kilometer east of the highway at this point. The elevation above sea
level is 7600-7700 feet, and the rolling mountains are covered with
pine-oak forest, with grassy areas in the valleys.
At the time of our visit (March 26-29, 1959), these Colias were
in full flight, and almost all specimens were fresh. The males were far
more numerous than the females, but as every female seen was usually
collected, the data given in Table 1 on relative abundance of the two
female color phases may be considered fairly accurate considering the
small number involved (for these dates). The frequency of white
females in this locality is within the range of frequency (10 to 20%)
found in the mountains of the western United States (Hovanitz,
1950b).
TABLE 1. Frequenqr of color phases of female Colias philodice m central
Chiapas, Mexico.
Yellow Females White Females Total Per cent White ± S.£.
11 2 13 15.38±10.0%
LITERATURE CITED
HOVANITZ, W. 1950a. The biology of Colias butterflies.
I. The distribution of the North American species. Wasmann Biol.
8: 49-75.
HOVANITZ, W, 1950b, The biology of Colias butterflies.
II, Parallel geographical variation of dimorphic color phases in North
American species. Wasmann J. Biol. 8: 197-219.
194
Journal of Research on the Lepiduptera 1 (3) :195-200, 1963
1140 \V'\ Orange Grove Aie., Arcadia, California, U.S.A.
© Copyright 1^6}
NOTES ON THE EARLY STAGES OF
TWO CALIFORNIA GEOMETRIDS
JOHN ADAMS COMSTOCK
Del Mar, California
AETHALOIDA PACKARDARIA
This moth was described by Hulst in 1888, as Hemerophila
packardafia. He subsequently created two synonyms for the same insect,
namely, Selidosema lachrymosa (1898, described from Los Angeles
County), and Selidosema homopteroides (1901, from Oregon).
Aethaloida packardaria was pictured by McDunnough in 1920, on
Plate VII, fig. ii of his Cleorini Studies.
In 1937 I published a description and illustration of the mature
larva, from an example taken in Bouquet Canyon, Los Angeles County.
This year ( 1962) I was able to obtain eggs and rear the species through
to maturity, which will serve to supplement and amplify the former
incomplete record.
Eggs were secured June 27 from captive females. Ceanothus and
Adenostoma were supplied to them, and oviposition occurred in clusters
on the tips of twigs. It was noted that the bases of the eggs in each
cluster pointed outward.
EGG: 0.8 mm. high, x 0.6 mm. at the widest point. The form is
subovoid, the base wider and the top tapering to a well rounded and
smooth surface, with no discernible micropyle. The base tapers inward
at the circumference and bears a circlet of pearl-like protrusions, about
20 in number, enclosing a granular depressed floor.
The surface of the egg above the circlet is finely granular, and does
not bear ridges. The color, when first laid, is dull olive-green. Later it
becomes darker.
Eggs laid June 27 hatched July 2 and 3, 1962. The illustration
(fig. lA) , shows the egg in lateral aspect.
FIRST INSTAR LARVA: Length, 2 mm. Width of a typical
body segment approximately 0.15 mm.
Head; width, 0.35 mm. Color, jet black except for a narrow trans-
verse bar on the labrum, which is dull white. The ocelli are black, and
the setae white.
The body is predominantly black. The thoracic segments taper from
the widest next to the head to the third which grades into the 0.15
195
196
COMSTOCK
/. Res. Lepid.
mm. wide first abdominal segment. The dorsum is solidly black. Laterally
on each segment there is a large subtriangulate white spot on each side,
which extends up over the dorso-lateral edge.
The legs are black, as are also the prolegs. There are rows of short
white setae, the most prominent of which is in longitudinal alignment
above the white spots.
The first instar larva is shown on figure IB. No attempt has been
made to accurately map the setae, but preserved larvae are available
for this purpose.
SECOND INSTAR LARVA, observed July 7, ’62. Length, 4.5 mm.
Body width, 0.4 mm., as is also the head width.
Head, black, as in the first instar.
The body is a dull maroon over the dorsum. The white triangulate
spots along the side persist as clumps of round spots in groups of ap-
proximately three, close together, or fusing with each other. There is a
transverse line of round white dots on the front margin of the first
segment.
The venter is dull maroon.
This batch of larvae did not thrive well, probably as a result of
too quick drying of the food plant. Mature larvae were later beaten
from Adeno stoma and Ceanothus, and were carried through to maturity.
My previous published description and illustration of the full grown
larvae (1937) does not need to be repeated, but a drawing is here
included on figure 1C.
PUPA: Length, 13.5 mm. Greatest width, 3 mm. Color, black on
wing cases, thoracic and head portions; red -brown on the abdominal
segments, with a darker shading on the segmental juncmres, and the
cauda.
Two black knobs protrude from the front of the head. The maxillae
and antennae reach to the wing margins. The cremaster consists of a
knobbed triangulate black element, from which arises a pair of rela-
tively long recurved spikes, and along the sides, three short red-brown
recurved booklets. The spiracles are concolorous with the body, and
relatively inconspicuous. The pupa is pictured on figure ID.
BIBLIOGRAPHY
COMSTOCK, JOHN ADAMS. 1937. Miscellaneous Notes on Western
Lepidoptera. Aethaloida packmdaria. Bull. So. Calif. Acad. Sci. 36 (3) :
41-124. PI. 47.
HULST, G. D. 1888. Hemorophila packardaria. Ent. Amer. Ill: 217.
HULST, G. D. 1888. Hemerophila packardaria. Ent. Amer. 111:217.
. 1898. Selidosema lachrymosa. Can. Ent. 30: 194.
— . 1901. Selidosema homopteroides. Jour. N. Y. Ent. Soc.
8: 219.
McDUNNOUGH, J. H. 1920. Studies in North American Cleorini, (Geo-
metridae) Bull. Dept, of Agric. Canada. 18: 37-38. Plate VII, fig. 11.
197
i(}):i9^-200, 196}
EARLY STAGES
Fig. 1, Early stages of Aethaloida packardaria (Hulst). A. Egg, lateral
aspect, enlarged X 50. B. First Instar larva, dorsal aspect, enlarged X 30,
C. Mature larva, lateral aspect, enlarged X 3. D. Pupa, ventral aspect,
enlarged X 5.
Reproduced from water color drawing hy the author.
PROCHOERODES FORFICARIA
In 1956 I published notes on the larva and pupa of the geometrid
moth, Prochoeroides forficaria (Guenee) from larvae obtained in Gila
County, Arizona.
198
COMSTOCK
/. R<s. Lcpid.
Dr. McDunnough’s published comments on this species (1940)
suggest that the Arizona specimens from which I described the larva
and pupa are closer to form catenulata Grote, and that this "could very
readily be considered a good species.”
I have recently reared typical P. forficaria from material taken at
Del Mar, and the following notes, with illustrations, give a more
accurate analysis of the species designated
Prochoerodes forficaria
Eggs were laid April 10, 1962 in a rearing jar. They were laid on
their sides, some singly, others in small bunches.
EGG: When first laid, bright green. Later they became dark, and
prior to hatching were black. The form was an elongate oval, with about
16 ridges extending from base to micropyle. There were no horizontal
ridges or lines. The micropyle was pitted.
The average length of the egg was 0.9 mm. and the width, 0.5 mm.
There was considerable variation in size and form, some examples
being elongate and somewhat less rounded at the ends.
The eggs hatched April 21 and 22, the larvae usually emerging
from the side, leaving the remainder of the shell intact.
Not knowing the food plant, the following were tried; lettuce,
dandelion, plantain, grass, fuschia, Ceanothus and Cneridium, none of
which proved acceptable. They finally accepted willow.
EIRST INSTAR LARVA: Length, 4. mm. Width, 0.2 mm. Head
width, 0.4 mm. The head is relatively large, and is mottled black and
brown. Ocelli, black.
The body is cylindrical, elongate, and very narrow. The ground
color is slaty gray. There is a broad longitudinal black band on the
dorsum, with occasional darker spots on some of the segments. The
lower lateral surface is a translucent gray-straw. The entire ventral
surface is black. The single pair of prolegs, and anal prolegs, are dark
on the lateral surfaces and lighter on the ventral surfaces. The setae are
white, and very minute.
The first instar larva is illustrated on figure 2B.
In this phase the larvae were exceedingly active, dropping on a
thread when touched.
Larva of 11 mm. in length, — head width, 1.55 mm.
The body is cylindrical, its entire surface being mottled red-brown,
on the dorsum there are indefinite quadrate patches of darker brown,
more clearly defined in the thoracic and caudal areas, but nowhere
connected sufficiently to form an unbroken band. The true legs are
yellow-brown, and the prolegs concolorous with the body. The latter
are prominent and conspicuous.
MATURE LARVA: Noted on May 5, at the beginning of the
final instar. Length of larva, 22 mm. Head width, 2.4 mm.
The body color is light wood brown, the head a shade lighter. The
body is mottled with darker spots and dashes, and its surface is crossed
transversely by numerous fine creases and ridges.
Fig. 2. Egg, larva and pupa of Prochoerodes forficaria. A. Egg, lateral
aspect, enlarged X 40. B. First instar larva, lateral aspect, enlarged X 24.
C. Mature larva, enlarged X 5. D. Pupa, ventral aspea, enlarged X 5.
E. Cremaster, highly magnified.
Reproduced from water color drawing by the author.
200
COMSTOCK
/. Res. Lepid.
Middorsally there is an indefinite longitudinal band of brown,
nowhere clearly defined. Spiracularly there is a pinkish irregular band,
enlarging around each spiracle, and contracting on the segmental junc-
tures. Near each spiracle there are two or three round dots. The legs
are slightly translucent, and the prolegs are concolorous with the body,
or slightly tinged with yellow.
There are numerous small papillae scattered over the body, two
pairs of which in the dorso-caudal area are slightly larger. All papillae
bear short yellow-brown setae. This is the larval phase that is pictured
on figure 2C.
On May 29, the larva measured 30 mm., the head width being 2.4
mm. as before. The chief difference in this prepupal stage is that there
is less definite ornamentation, although the pinkish area following the
spiracles is clearly defined, and the spiracles are more conspicuous, with
dark rims and a surrounding rim of yellow.
In the final phase the larva becomes sluggish, and spends much
time resting on a twig, or the midrib of a leaf, where it is ideally camou-
flaged. The first example pupated June 9, 1962.
PUPA: Length, 17 mm. Width through middle portion, 5 mm.
The color is brown, with the eyes and terminal segments darker.
The antennae extend slightly beyond the wing margins. The wing
cases and appendanges are overlaid with fine transverse brown lines
and dashes. The dorsal surface, and the abdominal segments are heavily
spotted with dark brown and black. The spiracles are concolorous with
the body and are inconspicuous.
The cremaster is formed by a double ovoid extension, narrowing
down to a clubbed shaft, which ends in two recurved tips that are arched
latero-ventrally. Near the base of the shaft there are six small yellow-
brown booklets, three on each side. These arch medially. The pupa is
illustrated on figure 2D and an enlarged figure of the cremaster on
figure 2E.
The first imago emerged June 21, 1962.
In 1926 Stanley E. Flanders recorded the recovery of three species
of parasites from larvae of Prochoerodes foriicaria, namely, Amblyteles
caeruleus (Cr.), A. zebratus (Cr.), and Euplectrus sp.
We have very little information on the range of P. forficaria but
Dr, McDunnough’s description of the southern British Columbia form
combinata suggests that our typical species ranges northward through
Oregon and perhaps Washington, where it grades into combinata.
We still need information as to its southern and eastern range.
BIBLIOGRAPHY
COMSTOCK, JOHN ADAMS. 1956. Scientific Notes. Prochoerodes for-
ficaria Gn. Larva, Pupa. Bull. So. Calif. Acad. Sci. 55 (3) : pp. 180-181.
FLANDERS, STANLEY E. 1926. Notes on Parasites at Saticoy, California
During the Year 1925. Pan. Pac. Ent. 2 (7): pp. 157-158.
McDUNNOUGH, J. H. 1940. New North American Geometridae with
Notes. 111. Can. Ent. 72(5): pp. 90-103.
Journal of Research on the Lepidoptera 1 ( 3 ) :201-208, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright
GEOGRAPHICAL DISTRIBUTION AND VARIATION
OF THE GENUS ARGYNNIS
III. ARGYNNIS DIANA
WILLIAM HOVANITZ
California Arboretum Foundation, Inc., Arcadia, California, and
Los Angeles State College, Los Angeles, California
The Diana fritillary, Argynnis diana Cramer, departs from
the most usual color pattern for the genus as the pattern is developed
in most species of the Nearctic and the Palearctic. The male and female
both have the basal two-thirds of the wings a brownish black. This
distribution of pigment is in part the reverse of the distribution for
Argynnis idalia, in which the black pigment is primarily restricted to
the outer half of the wings. It has already been pointed out (Hovanitz,
1963) that the distribution of this pigment for idalia is diametrically
opposed to the ordinary tendencies in other members of the genus in
which the black pigment is normally distributed in the area of the
basal regions. This is the pattern arrangement for Argynnis diana, A.
cyhele, A. leto, A. nokomis and others in North America which show an
extension of basal melanism. The remainder of the wings shows the
usual pattern arrangement of black dots and black striping which is
typical of the genus Argynnis in any part of the world. In the male,
however, these black markings are greatly diminshed and restricted in
most cases, especially on the hind wing, so that the outer one-third of
the wings on the upper side are nearly a solid brownish-orange color.
On the female this restriction of the black does not occur. It is in fact
greatly extended so that not only is the outer margin all black, but the
black-veined stripes are wide and extend from the basal black to the
margin, and the black dots are large and converge to form black stripes
running nearly from the anal margin to the inner margin at least on
the fore wing. The part of the wings then remaining that would nor-
mally be a brown color in Argynnis is an irridescent blue. On the face
of it, the female diana is a black butterfly with marginal and sub-
marginal rows of blue spots. This dimorphism of the sexes of A. diana
is the greatest expression in North American Argynnis of this type of
variation. It is, however, nearly duplicated by some races of Argynnis
nokomis, to which it may be closely related (fig. 1 ) .
The under side of the wings also shows a very different aspect than
the usual Argynnis pattern. This alteration of the pattern is more than
the usual general suffusion of color over the wing but effects the pattern
201
202
HOVANITZ
/. Res. Lcpid.
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Fig. L Argynnis diana, upper side. Top, male; bottom, female. The
blue is irridescent and may show only in certain directions of certain quality
of light. ( See fig. 2 ) .
I (}):20I-2o8, 1963
ARGYNNIS DISTRIBUTION
203
Fig. 2. Same as fig. 1, only under side. $ Salem, Roanoke Co., Vir-
ginia. June 13, 1937. C. W. Gottschalk. 2 Montgomery Co., Virginia. Aug.
16, 1902. E. A, Smyth, Jr.
204
HOVANITZ
/. Res. Lepid.
elements in such a way as to be most certainly of fundamental embryo-
logical nature. By this is meant the almost complete obliteration of the
pattern elements on the underside of the hindwings in both the male
and the female, with a slight degree of the same on the fore wings. In
this development, A. diana is also most directly opposed to A. idalia in
which the spots and pattern elements are well developed, even more so
than in many other Argynnis. The pattern observed on the diana hind
wing underside looks as though all markings had been wiped clean
between the most marginal row of small spots and the usual Argynnis
row of large silvery spots, substituting therefore, a uniform bluish
black on the female, light brown in the male; likewise for the area
from the base of the wing outward to the same row of spots nearly all
markings of the usual Argynnis type appear to have been wiped clean
except for a few minor points and substituting therefore a brownish-
black on the female and light brown in the male. A small part of the
central row of silvery spots of the usual Argynnis pattern remains, along
with a small part of the cell spot and most marginal row in both the
male and the female. The inner two-thirds of the fore wings on the
under side in both sexes are similar to the usual Argynnis pattern.
Interestingly enough, there is blue irridescence in this part of the wings
of the male, even though the upper side of the male wings do not
show it. ( fig. 2 )
A similar development of the pattern elements on the under side of
Argynnis wings occurs in the groups related to A. paphia, A. pandora,
A. laodice, A. childreni, etc. of Eurasia. These are the groups which
have a distributional range through areas of higher rainfall, and thus
more humid air, combined with higher summer temperatures. Some
membrs of these groups also have a development of sexual dimorphism
similar to that of A. diana and A. nokomis, even to the extent of having
blue or bluish females while the male is the usual orange-brown color.
More complete correlations between wing color, pattern type and insect
size with climatic differences in various parts of the distributional range
of Argynnis will come in a later issue.
The geographical distribution of Argynnis diana is a rather re-
stricted one (fig. 3), hardly being more than 10° from the most
northern part of its range to the most southern part and 15° from the
most easterly to the most westerly part of its range. This area in miles
is abou 1000 miles from east to west and three to four hundred miles
from north to south. Actually, the present known distributional range
is shaped somewhat like a triangle, with an acute angle in the far west
in Arkansas. Except for a collection in western Pennsylvania which
may be inaccurate, the species is known only as far north as southern
West Virginia and central Virginia. The most southern localities are
southern South Carolina and northern Georgia and an area of north-
central Arkansas. There are records of the species from southern
Illinois. It is likely that they may be found in southern Missouri, southern
(});20i-2o8, 1^6}
ARGYNNIS DISTRIBUTION
205
NORTH AMERICA
Fig. 3. Map showing the North American distribution of Argynnis
diana and A. nokomis. Note the wide expanse of isolation between these two
closely related butterflies; also the wide isolation between parts of the range
of A. nokomis. Details on the geographical variation of nokomis will come in
the next number in this series.
206
HOVANITZ
/. Res. Lepid.
Indiana and southern Ohio in addition, though I have not seen specimens
from these locations.
This species, despite its striking colors and size, is a relatively rare
species due to the limited areas in which it lives. There is no evidence
that at any time in the past the species was any less local than it is now,
even though it is said that the deforestation of many parts of the east
have contributed to its demise in many areas (Clark, 1951). The
species exists in parts of North America away from the most heavily
inhabited regions and therefore collectors have not had as much experi-
ence with it as with other species. Undoubtedly, the restricted habitat
of the larvae contributes most to its own restricted range and the factors
that are essential for larval environment are not known. Judging how-
ever from the habitats preferred by the adults and especially the females
which is usually highly indicative, the species finds its home in areas
of cold seepages and streams on the coastal plain, or in deep, damp,
well wooded valleys or ravines, or on damp wooded mountain sides
where there are cold streams (Clark, 1951). This corresponds to the
habitat of A. nokomis, the western related species (fig. 1), which
occurs in dry desert regions but only along cold streams where there
is an abundance of boggy vegetation and locally a "wet-tropical” micro-
habitat as at Round Valley, Inyo County, California. This type of
habitat is very local, not only in the desert regions of the west but also
apparently in the east and is probably more responsible for the restric-
tion of range of diana than the removal of the forest cover over exten-
sive areas of the eastern parts of North America.
Clark (1951) gives the most complete description of the distribu-
tion, variation and habits of this species than has ever been given. The
following is from his description :
"The first males to appear are smaller than those that emerge later,
and the males from Highland County and from the higher latitudes
in the mountains farther south are always small, resembling the earliest
males from other regions.
"On the Coastal plain the males occasionally have on the under side
of the hind wings at the end of the cell a conspicuous silver spot
bordered inwardly and outwardly by black lines, corresponding to the
silver spot in the same position in S. cybele though smaller, seldom
reaching more than halfway across the cell, and the silver markings on
the outer portion of the hind wings may be enlarged. Occasionally on
the upper surface of the fore wings there are broad light dashes in the
black ground color beyond the cell in the interspaces between veins 3
and 4, 4 and 5, 5 and 6, corresponding to the light dashes on the under
side though with indefinite borders. So far as we have seen, these
features are confined to males from the Coastal Virginia Plain. The
majority of the males from the Coastal Plain, however, do not differ
in any way from others from the mountains.
"Dr. Henry Skinner has pointed out that the males differ materially
l(})t20l-20i, 1^6}
ARGYNNIS DISTRIBUTION
207
in the number and size of the black spots on the upper surface of the
hind wings. We have seen males, both from the Coastal Plain and the
mountains, with the light border of the hind wings almost immaculate.
There is some variations in the size of the black spots on the fore wings,
and veins 2, 3, and 4, may be narrowly or rather broadly infuscated.
The contrast between the dark basal and light outer portions of the
under surface of the hind wings is sometimes accentuated. These varia-
tions seem to bear no relation to locality,
"Dr. Skinner said that females from eastern Tennessee, western
North Carolina, and southern Illinois are larger than those found in
Virginia; but the females from the lower altitudes in southwestern
Virginia appear to be quite as large as any from farther south. Dr.
Skinner pointed out that the females vary a great deal in the degree of
silver beneath and also in the band of large bluish or greenish spots
on the hind wings above. In sorhe specimens these spots are large, and
in others they are confined to a small area around the black spots.
The number and size of the cream-colored or white spots on the upper
side of the fore wings is also quite variable.”
It appears from my own observations as well as from those of others
that there is quite a lot of variability in this species; most of these
variations however appear to be individual rather than population in
scope, with the exception of those that are directly related to the eleva-
tion differences described by Clark. Occasionally the blue irridescence
has been changed to green. Statistical study of large samples seems to be
called for to bring this variation out of the realm of guesswork.
The species flies over a long season, males coming out in the middle
of June and females slightly later with a height of emergence in the
middle of August and extending into September.
Most authors in discussing this species in handbooks comment on
mimicry between this species and some other species which inhabit the
same general geographical area. Ehrlich (1961) states "The female
presumably mimics the distasteful Aristolochia swallowtail, Battus
philenor,” Klots (1951) states "This is cited as a case of mimicry, the
supposed model being the black and blue Papilio philenor, a swallowtail
supposedly distasteful to birds.” Scudder (1889) considers the female
is a case of parastatic mimicry with Basilarchia aslyanax. It is fortunate
that most authors leave the case in doubt as most all situations of this
sort are wild grasps at straws in the wind. The relationships in appear-
ance of many butterflies in the same general geographical area, such as
these three, A, diana in the east and A. nokomis in the southwest with
Papilio philenof and Basilarchia astyanax are exceedingly interesting
and deserve some intensive study. Too often these studies are superficial
and involve some transference of a man’s way of thinking to a butterfly
or a bird. As purely a suggestion, it appears that a more satisfactory
solution may be found in the physiological relationship of the butterfly
with its physical environment, in which the colors are related in some
208
HOVANITZ
/. Res. Lepid.
way to light reflection from the butterfly’s wings (U.V. light perhaps)
which reflections are developed in the butterflies living in a warmer
or more humid environment.
LITERATURE CITED
CLARK, A. H. 1951. The Butterflies of Virginia. Smithsonian Misc.
Collections, 116(7): 1-95.
EHRLICH, PAUL and ANNE H. 1961. How to know the butterflies.
Brown, Dubuque, Iowa.
HOVANITZ, WILLIAM. 1963. Geographical distribution and variation
of the genus Argynnis. J. Res. Lepid. 1(2): 117-123.
KLOTS, A. H. 195 L Field guide to the butterflies. Houghton Mifflin,
Boston.
SCUDDER, S. 1889. The Butterflies of the Eastern United States. Cam-
bridge, Mass.
Journal of Research on the Lepidoptem 1 (3 ) :209-221, 1963
1140 W, Orange Grove Ave., Arcadia, California, U.S,A.
© Copyright 196}
THE GENERIC, SPECIFIC AND LOWER CATEGORY
NAMES OF THE NEARCTIC BUTTERFLIES
PART 2 — The Genus CoUas
PADDY. McHENRY
1 032 E. Santa Amta^ Burbank, California
The names included io this genus are presented in the same
manner as were those of the genus Pieris in the first part of this series
{]om. Res. Lepid. 1(1) :63-71, 1962).
W, Hovanitz has separated the members of the genus into specific
groups as best he felt was possible in ¥iew of the extensive hybridiza-
tion in the genus. ‘
Tlie genus Zerem Hiibner, although considered a subgenus of
Colias by some students, will be dealt with separately in a subsequent
part.
The data for chfysojheme Esper has been listed under the eurytheme
group purely as a convenient place in which to give this reference;
should a relationship be established between chrysotheme and eurytheme,
the former must, of course, be accorded priority.
The name notatus has been presented in the philodice group under
the authorship of Megerk and also under the authorship of Clark and
Clark because of its present uncertain nomenclatorial status.
LIST OF GENERIC NAMES USED OR AVAILABLE FOR
COLIAS,
COLIAS Fabricus.
Type: hyde Linnaeus.
EURYMUS Horsfield.
Type: h'yde Linnaeus.
COLIAS FABRICIUS. 1807 [before 19 Deep Io K, Illiger. Magazin fur-
Insekteokunde 6: 284, oo. 24, Includes among others ^Pap . . .
Hyde’* and Rhamni" both considered to be of Linnaeus, 1758.
Type. P[apiUo\ D^anms]. {Candidas'] hyde Linnaeus. 1758. Syst. Nat.
10th. Ed. 1: 469, no. 71.
Type Selection. 30 Sept. 1943. Opinions & Declarations Rendered by
the International Commission on Zoological Nomenclature 2(13):
109-121, The type is declared to be hyde Linnaeus, 1758. This
declaration invalidates the selection of ^Colias Rhamni, Fab,” (con-
sidered to be rhamni Linnaeus, 1758) by Latreille, 1810, Consid.
Gen. Anim. Crust. Arach. Ins, : 440.
EURYMUS HORSFIELD, (Swainson Ms. name). 1829. [24 June]^.
Catal. Lepid. Ins. Contained Mus. Honor. East Indies Company
(2) : 134. Gives only ^^Colias Hyale” which is considered to be
hyale Linnaeus, 1758. The name is a homonym of Eurymas Rafin-
esque, 1815^.
Type. P[apilm]. D\anaus]. [Candidm] hyale Linnaeus. 1758. Syst. Nat.
10th. Ed. 1 : 469, oo. 71.
Type Selection. Horsfield. 1829 [24 June]^. Ibid. (2) : 134. "Mr. Swain-
son gives Colias Hyale as the type of this subgenus . .
209
210
McHENRY
/. Res. Lepid.
LIST OF SPECIES AND LOWER CATEGORY NAMES USED OR
AVAILABLE FOR COLIAS
1. COLIAS BEHRII W. H.
EDWARDS.
behrii W. H. Edwards.
ccmescens (J. A. Comstock).
2. COLIAS BOOTHII CURTIS.
boothii Curtis.
chione Curtis.
3. COLIAS EURYTHEME
BOISDUVAL.
alba Strecker.
alba Strecker.
amphidusa Boisduval.
ariadne W. H. Edwards.
californiana Menetries
chrysotheme (Esper).
eurytheme Boisduval.
flam Strecker.
fumosa Strecker.
intermedia Cockerell.
keewaydin W. H. Edwards.
pallida Cockerell.
pallida Cockerell.
pallida Cockerell.
rudkini (Gunder).
unicitrina (Gunder).
4. COLIAS GIGANTEA
STRECKER.
gigantea Strecker.
harroweri Klots.
marjorie Chermock & Chermock.
ma^i Chermock & Chermock.
pelidneides Staudinger.
5. COLIAS HECLA LEFEBVRE,
chrysothemoides Verity.
glacialis M’Lachlan.
hecla Lefebvre.
hela Strecker.
palamedes Hemming.
pallida Skinner.
6. COLIAS MEADII W. H.
EDWARDS.
elis Strecker.
meadii W. H. Edwards.
medi (Gunder).
7. COLIAS NASTES
BOISDUVAL.
alias ka Bang-Haas.
cocandicides Verity.
gueneei Avinoff.,
harperi (Gunder).
moina Strecker.
nastes Boisduval.
ob scut at a Verity.
palliflava (McDunnough) .
rossii Guenee.
streckeri Grum-Grshimailo.
subarctica (McDunnough) .
thula Hovanitz.
8. COLIAS OCCIDENTALIS
SCUDDER.
alba Strecker.
alberta Bowman.
alexandra W. H. Edwards.
astraea W. H. Edwards.
bar bar a Henry Edwards.
Christina W. H. Edwards.
chrysomelas Henry Edwards.
edtvardsii W. H. Edwards.
emilia W. H. Edwards.
harfordii Henry Edwards.
hatui (Barnes & Benjamin).
krauthii Klots.
lambillioni Dufrane.
martini (Gunder).
occidentalis Scudder.
pallida Cockerell.
pallida Cockerell.
pallidissima Bowman,
shastae (Barnes & Benjamin).
weaverae ( Gunder ) .
9. COLIAS PALAENO
(LINNAEUS).
chippewa W. H. Edwards.
helena W. H. Edwards.
kohlsaati (Gunder).
lapponica Staudinger.
palaeno (Linnaeus).
philomene (Hubner).
werdandi Herrich-Schaffer.
10. COLIAS PELIDNE
BOISDUVAL & LECONTE.
intertor Scudder.
isni (Barnes & Benjamin).
labradorensis Scudder.
laurentina (Scudder).
minisni Bean.
mira Verity.
moeschleri Grum-Grshimailo.
nepi (Barnes & Benjamin).
neri (Barnes & Benjamin).
pelidne Boisduval & LeConte.
skinneri Barnes.
solivaga W. H. Edwards.
1 1 . COLIAS PHILODICE
GODART.
alba Strecker.
alba F. Chermock.
albida (F. Chermock),
albinic W. H. Edwards.
ant hy ale (Hubner).
(});209~22l, 196}
NAMES — Pt. 2
211
autumndis Cockerell.
ehrmanni F. Chermock.
eriphyle W. H. Edwards.
europome ( Haworth ) .
hagenii W. H. Edwards.
hybrida Strecker.
inversata Nakahara.
kootenai Cockle.
laurae (F. Chermock).
lutetincta Wolcott.
melanic W. H. Edwards.
melanic W. H. Edwards,
minor F. Chermock.
miscidice (Scudder).
nig Strecker.
nigricosta (F. Chermock).
nigfidice ( Scudder ) .
nigrina Strecker,
nigrofasciata Reiff.
notatus (Mergerle).
notatus (Clark & Clark).
pdlidice ( Scudder) .
philodice Godart.
plicaduta Nakahara.
raritus (Gunder).
reducta Dufrane.
rothkei Reiff.
s errata (F. Chermock).
suffusa Cockerell.
virida Strecker.
vitahunda Hovanitz^
12. COLIAS SCUDDERII
REAKIRT.
flavotincta Cockerell.
ruckesi Klots.
scudderii Reakirt.
1. COLIAS BEHRII W. H. EDWARDS.
hehrii, Colias W. H. Edwards. Oct. 1866. Proc. Ent. Soc. Phila. 6
(pp. 201-208): 201. "From 2 $, I $, received from Dr. Behr,
and taken among the Yo Semite mountains at an elevation of about
10,000 feet above the sea.” No dates given. Spelled behri by J. A.
Comstock, 20 May 1925, Bull. Sou. Calif. Acad. Sci. 24(1): 3.
canescens, Eurymus behri J. A. Comstock. 20 May 1925. Bull. Sou.
Calif. Acad, Sci. 24(1) : 3. "Type [$], Tioga Pass, Yosemite, Calif.,
Aug. 4, 1922. Six paratypes [$ $]. Same place and date. In the
collection of the Southwest Museum.”
2. COLIAS BOOTHII CURTIS.
chione, Colias Curtis. 1835. In J. Ross. Append. Narr. 2nd Voy.
Search North-West Pass. (Nat, Hist. Sect.), pp. Ixvi-lxvii, no, 11;
plate A, fig. 6. d & $ described. ”... they were captured from
the I4th of July to the 13th of August, 1830, and on the 19th
were in a very wasted state; on the l4th of July, of the following
year, one Colias only was taken.” No locality nor series data given
here.
boothii, Colias Curtis, 1835. In J. Ross. Append. Narr. 2nd. Voy.
Search North-West Pass. (Nat. Hist. Sect.), pp. Ixv-lxvi, no. 10;
plate A, figs. 3-5. d & $ described. No locality, series data nor
dates given here. Spelled boothi by Verity, 1905-1911, Rhopal.
Palaearaica, p. 238.
3. COLIAS EURYTHEME BOISDUVAL.
alba, Colias chrysotheme eurytheme Strecker. 1878 [Sept. - Ort.]"^. Butt.
& Moths N. Amer., Compl. Syn. Catal., p. 83, no. 60b. Cites $
figures in W. H. Edwards, Butt. N. Amer. [I]: plate Colias III, figs.
5-6, 1869. No locality, series data nor dates given here.
alba, Colias chrysotheme keewaydin Strecker. 1878 [Sept. - Oct.]"^. Butt.
& Moths N. Amer., Compl. Syn. Catal., p. 83, no. 60e. Cites $
figures in W. H. Edwards. Butt. N. Amer. [1]: plate Colias IV, figs.
8-9, 1869. No locality, series data nor dates given here.
amphidusa, Colias Boisduval. 1852 [Aug.]^ Ann. Soc. Ent. France. 2nd.
Ser. 10(2): 286-287, no. 14. d & 9 described. "Du nord de la
Californie.” No series data nor dates given. Spelled amphidona
by W. H. Edwards, 1869, Butt. N. Amer. [1]: [45].
ariadne, Colias W. H. Edwards. Jan. 1870. Trans. Amer. Ent. Soc. 3(?)
sign. 2: 12-13. ”1 d, 1 9, from the collection of Dr. Behr, and
taken at Mokeluma Hills, California.” No dates given.
212
McHENRY
/. Res. Lepid.
Californian, Colias edusa Menetries. Dec. 1855. Enum. Corp. Anim.
Mus. Imper. Acad. Sci. Petropolitaoae. Class. Ins. Ordo Lepid. ( 1 ) :
80, under no. 253. "Ces caraaeres indiques ci dessus se reproduisent
constamment sur environs 6 males et 4 femelles, tous rapportes de
la Nouvelle California par notre zele voyageur Wosnesensky.” No
dates given.
chrysotheme, Dan[aus]. Can{didus\ Esper. [1781]*. Die Schmetterlinge
1(2) heft 3: 89-90, no. 131; plate 65, figs. 3-4. d & $ described.
No locality, series data nor dates given. Note: This name is included
here only for convenience.
eurytheme, Colias Boisduval 1852 [Aug.]*. Ann. Soc. Ent. France. 2nd.
Ser. 10(2): 286, no. 13. "Commune dans toute la Californie.
Elle habite aussi le Mexique et quelques parties des Etats-Unis.”
No sex, series data, nor dates given. Spelled enegthenu by W. H.
Edwards, 1872, in F. V. Hayden, Prel. Kept. U. S. Geol. Surv. Mont.
& Port. Adj. Terr. 5th. Kept. Progress, p. 466.
flava, Colias chrysotheme eurytheme Strecker. 1878 [Sept. - Oct.]"^. Butt.
& Moths. N. Amer., Compl. Syn. Catal., p. 83, no. 60c. 2 described.
"Mus. Strecker." No locality, series data nor dates given.
fumosa, Colias eurytheme Strecker. 1900 [Mar. 9th-13th]^. Lepid.,
Rhopal. & Heter., Indig. & Exot. Suppl. No, 3: 19. "One $,
Colorado . . No date given.
intermedia, Colias eurytheme Cockerell. Mar. 1888 [after 9th]®. West
Amer. Scientist 4(35): 41-43 (in pt.). 2 described. "In this
locality (Swift Creek, Custer Co., Colo[rado] , . No series data
nor dates given.
keewaydin, Colias W. H. Edwards. [Sept. 1869, on or before 13th]'^,
Butt. N. Amer. [I](4) : [47]-[49]; plate (15), figs. 1-9. d & 2
described. '^Found in the valley of the Mississippi from Nebraska
and Illionis to Texas and westward to the Pacific, occupying much
the same region as Eurytheme, but apparently less common and
more local than that species. Also occasionally found in the Middle
States and Canada." "There appear to be two broods during the
year, the insects being most abundant early in the spring, in faa,
before Eurytheme makes its appearance.” No series data nor dates
given.
pallida, Colias eurytheme Cockerell. Nov. 1887. West. Amer. Scientist
3(31): 217. 2 described. "... near West Cliff, Custer Co.,
Colorado.” No dates given.
pallida, Colias eurytheme intermedia Cockerell. Mar. 1888 [after 9th]®.
West. Amer. Scientist 4(35): 41-43 (in pt.), "... Whitish
females occur of . . . intermedia, these may be called pallida.” No
locality, series data nor dates given,
pallida, Colias eurytheme keewaydin Cockerell. Mar. 1888 [after 9th]®.
West. Amer. Scientist 4(35): 41-43 (in pt.). . . Whitish
females occur of . . . keewaydin . . . these may be called pallida.”
No locality, series data nor dates given.
fudkini, Eurymus eurytheme Gunder. 31 Dec. 1932. Can. Ent. 64(12) :
211 -21 k. "Holotype - 2 . . . San Marino, Los Angeles Co., Calif.,
May 23, 1931. Type in Author’s coil.”
unicitrina, Eurymus eurytheme amphidusa Gunder. 8 May 1924. Ent.
News 35(5): 158; plate 2, fig. J. "Holotype (Author’s Coll.)
Upland, Los Angeles County, California; August 2, 1921.”
4. COLIAS GIGANTEA STRECKER.
gigantea, Colias pelidne Strecker. 1900 [Mar. 9th"13th]^. Lepid.,
Rhopal. & Heter., Indig. & Exot. Suppl. No. 3: 19. "Five $ S,
1 2 2, west coast Hudson Bay, above Fort York, Archdeacon
Kirby.” No dates given.
t(}):ao$-22ig 1^6}
NAMES ™ Pt. 2
213
hatfoweri, Colias gigantea Klots. 22 Mar. 1940. Amer. Mus. Nov. No.
1054: 4-6. "Type Lot All are from Clear Creek, vicinity Lower
Green River Lake, Sublette Co., Wyo[ming]., alt. 8400 ft. Holotype
male, three male and two female paratypes, August 3, 1935 • .
allotype female, nine male and one female paratypes, July 16,
1939...”
marjofie, Colias Christina mayi Chermock & Chermock. 30 Apr. 1940.
Can. Ent. 72(4): 82. "Holotype - $, June 26, 1933, Riding
Mountains, Manitoba. Paratypes - 1-50, same locality.” Page 8N
"... holotypes will be deposited in Canadian National Collec-
tion . . .”
mayh Colias Christina Chermock & Chermock. 30 Apr. 1940. Can.
Ent. 72(4): 81. "Holotype - d, July 1, 1933, Riding Mountains,
Manitoba. Allotype - 2, July 1, 1933, same locality. Paratypes -
1 to 150, same locality.” Page 81: " . . . holotypes will be
deposited in the Canadian National Colleaion and the allotypes
in the Carnegie Museum.”
pelidneides, Colias palaeno Staudinger. May 1901. In Staudinger &
Rebel. Catal. Lepid. Palaearct. Faun. 3rd. Ed., p. 15, no. 86i. $
described. "Amer. bor. (Hudson Bay, Alaska).” No series data
nor dates given.
5. COLIAS HECLA LEFEBVRE.
chrysothemoides, Colias hecla Verity. 1905-1911. Rhopal. Palaearctica:
p. 356; p. xxxvii; plate LXXI, figs. 22-23; plate LXXI explanation
page, figs. 22-23. "22 . . .d (Territoire de Barren, 114°, 30’ long.
O., 67° 40’ lat. N., Am. boreale).” "23 . . .9 ([same as 22]).”
Collection of British Museum. No series data nor dates given.
glacialis, Colias hecla M’Lachlan. 23 May 1878. Jour. Linnean Soc.
(London), Zoology 14(74): 108. "Two $ and one $ from lat.
81° 45’, August 12th, 1876, and one 2 from Hales Sound, lat.
79° . . . also a much crippled $ . . . from Discovery Bay, July
18th, 1876 . . .”
hecUf Colias Lefebvre. 1836 [after 6 Apr.]’°. Ann. Soc. Ent. France
5(?)': 383-387; plate 9, figs. 3-6. ^ & 2 described. " V
Islande est la patrie . . .”. No series data nor dates given.
hela, Colias Strecker. Sept. 1880. Bull. Brooklyn Ent. Soc. 3(5) : 33-34.
"... the present descriptions are from 1 d and 2 2 2 ; a few more
were taken, but these are all I have had the opportunity of exam-
ining. They were captured ... a considerable distance above Fort
Churchill on west coast of Hudson’s Bay; . . .”. No dates given.
palamedes, Colias hecla Hemming. May 1934. Stylops, Jour. axon.
Ent. (5) : 98, no. 17. A new name for Colias hecla pallida
Skinner which he states is a homonym of Colias erato pallida
Staudinger (1861, in Staud. & Wocke, Cat. Lepid. Europ., p. 3).
pallida, Colias hecla Skinner (Skinner & Mengel Ms. name). 1 Mar.
1892. Ent. News 3(3) : 49; plate 2, fig. 4. 2 figured. "... from
Greenland.” No date given, A homonym of Colias erato pallida
Staudinger (1861, in Staud. & Wocke, Cat. Lepid. Europ., p. 3).
Spelled palida by Holland, 1931, Butt. Book, rev, ed., p. 298.
6. COLIAS MEADII W. H. EDWARDS.
elis, Colias Strecker. 10 Mar. 1885. Proc. Acad. Nat. Sci. Phila. [37]
(sign. 2) : 24. Description concluded on page 25 (sign. 3) : 7 Apr.
1885. "Taken ... at an elevation of 10,000 feet, on the summit
of "Kicking Horse Pass,” in the Rocky Mountains, between Alberta
Territory and British Columbia, at the boundary between the
United States and the British possessions, about 300 miles north
of Montana.” "Capt. Geddes took about fifteen examples, all
females ...” No dates given.
214
McHENRY
/. Res. Lcpid.
meadii, Colias W. H. Edwards. Feb. 1871. Trans. Amer. Ent. Soc.
3(.^) sign. 34: 267-268. 6 & $ described. No locality, series
data nor dates given. In his Butt. N. Amer. [1]: 59, 1868-1872,
he says: "Taken in Colorado ... in the month of July, 1871.” He
speaks of Mr. T. L. Mead taking 12 specimens on the "divide,” 34
specimens on "Mt. Lincoln” and 1 specimen at "Kenosha House.”
Spelled meadi by Holland, 1931, Butt. Book, rev. ed., p. 297.
medi, Eurymus meadii Gunder. 30 June 1934. Can. Ent. 66(6): 125.
"Holotype - $, . . . Beckenridge Peak (11,000 ft.), Empire County,
Colorado, Aug. 8, 1919. Type in Author’s coll.”
7. COLIAS NASTES BOISDUVAL.
aliaska, Colias nasPes Bang-Haas. 15 Aug. 1927. Horae Macrolepid.
Reg. Palaearctica 1(3) : 41. Figs. 24-25 on plate 5 were published
later. "Habitat: Alaska: Ramport 8. Juli, ...,3d 2 $ Spelled
alaskae by McDunnough, 1938, Mem. Sou. Calif. Acad. Sci. 1: 8,
no. 52e.
cocandicides, Colias nastes rossii Verity. 1905-1911. Rhopal. Palaearctica;
p. 355 (as rossii); p. xxxvii (as cocandicides); plate LXXI, figs. 7-9;
plate LXXI explanation page, figs. 7-9 (as rossii). "7 . . . d
(Territoire de Barren, 114°, 30’ long. 0., 67° 40’ lat. N., Am.
Iwreale)”. "8 . . . $ ([same as 7])”. "9 . . . d ... ([same as 7
& 8]) ”. No series data nor dates given.
gueneei, Colias nastes Avinoff. 10 Dec. 1935. In Holland & Avinoff.
Mem. Carnegie Mus. 12(2): sect. 5, sub-sect. 2: 13-14, no. 10;
& page 32; plate 27, figs. 1, 11, 21-22, 25-27, 31-32. d & $
described. Locality: Southampton Island. No series data nor dates
given.
harperi, Eurymus nastes moina Gunder, 31 Dec. 1932. Can. Ent. 64(12) :
278. "Holotype - 2 , . . .; Ft. Churchill, Manitoba, Can[ada]., July
19, 1932. Type in Author’s coll.”
moina, Colias Strecker. Sept. 1880. Bull. Brooklyn Ent. Soc. 3(5): 34.
d & 2 described. "Described from a large number of examples
mostly males.” ["a considerable distance above Fort Churchill on
West Coast of Hudson’s Bay” . . .] (taken from Strecker’s text
of hela, p. 34) .
nastes, Colias Boisduval. [1832]”. Icon. Hist. Lepid. Nouv. Peu Connu
(?): plate 8, figs. 4-5 (d). Pages 245-246 published later
( d & 2 ) . "Elle se trouve au Cap-Nord et en Islande. M. Sommer
m’en a aussi envoye plusieurs individus pris au Labrador.” No series
data nor dates given.
obscurata, Colias nastes streckeri Yenty. 1905-1911. Rhopal. Palaearctica;
pp. 354-355 (in pt.); p. xxxvi; plate LXXI, fig. 6; plate LXXI
explanation page, fig. 6. "6 . . . d ... (Lake Louise, Alberta)”.
No date given,
palliflava, Eurymus nastes streckeri McDunnough. 25 July 1927. Can.
Ent. 59(7): 153-154. d described. "... higher slopes of Mt.
McLean (above 6500 ft.) on July 12 . . .”. ". . . type No. 2425
in the Canadian National Collection, Ottawa”.
rossii, Colias nastes Guenee. 12 Oa. 1864. Ann. Soc. Ent. France.
4th. Ser. 4(2) : 199-200. d & 2 described. "... trois individus , .”.
No locality nor dates given here. Spelled rossi by Holland, 1931,
Butt. Book, rev. ed., p. 299.
streckeri, Colias nastes Grum-Grshimailo. 30 Apr. 1895. Horae Soc.
Ent. Rossicae 29 (1-2) : 290-291, no. 2. d & 2 described. "Specimen
unum hujus varietatis sub nomine ^'Colias Behrii?” a lepidopterologo
germanico D-re 0. Staudinger anno 1891, quattuor specimina, in
provincia Alberta ad Laggan collecta, a lepidopterologo americano
Dom. H. Strecker, cujus in honorem hanc forman nominavi, accepi”.
I (}) ‘.20^-221, 1^6}
NAMES --- Pt. 2
215
subarctica, Eurymus nastes McDunnough. 1 Dec. 1928. Cao. Ent.
60(11): 270^271. "Holotype. ^ Bernard Harbour, N. W. T.,
Aug. 9, 1915, . . . , No. 2863 in the National Collection, Ottawa.
Allotype. - 2 , same data. Paratypes. - 4 d , 4 $ , from same locality,
taken on various dates early in August”.
thula^ Colias nastes Hovanitz. [1955, Mar. 9-11 Oct,]’^. Wasmann
Jour. Biology 13(1): 1-8; figs. 1-2, "These collections [made in
1952 and 1953] of Colias consisted primarily of the species C. nastes
(thirty-five specimens) . . "The specimens with which we are
concerned here were collected at a camp located along this river
[Meade River, northwestern part of Alaska Territory] at a point
North 70° 45’ by 156° 30’ West”. "Holotype male and Allotype
female. Locality: Near the Mead River, Alaska Territory, Alaska,
N 70° 45’ X 156° 30’ W. Date: Male July 13, 1952, and female
July 11, 1952 . . . Depository: U . . . S . . . Nat. . . Mus. . Wash-
ington, D. C. Paratypes: One pair in the author’s collection at the
California Academy of Sciences, San Francisco, California, One pair
each in the Canadian National Collection at the Department of
Agriculture, Ottawa; the American Museum of Natural History,
New York; and the British Museum of Natural History, London.
These may be dated 1952 or 1953 . . . The remainder of the series
are io the collection of the Allan Hancock Foundation”.
8, COLIAS OCCIDENTALIS SCUDDER.
alha, Colias dexandra Strecker. 1878 [Sept. - Oct.]"^. Butt. & Moths N.
Amer., CompL Syo. Catal., p. 81, no. 55a. "... a white $ form
of rare occurence”. No. locality, series data nor dates given.
alberta, Colias eurytheme Bowman. 9 Mar, 1942. Can. Ent. 74(2) : 25.
"Holotype - 6, Wembley, Al[ber]ta,, June 24, 1925, in .the author’s
collection. Allotype - 2, Wembley, Alta., June 25, 1925, in the
author’s collection. Paratypes - 7 d , 2 2 , Wembley, Alta., June
14-25, 1925; 1 6,1 2, Beaver Lodge, Alta., June 29 and July 12,
1924; 1 6, Fort Vermilion, Alta., June 13, 1925; 1 2, Boswell,
B. C, July 22, 1927; ^2 2, Rolla, B. C, July 1 and 21, 1927.
Paratypes in the Canadian National Collection and in the collection
of the author.”
dexandra, Colias W. H. Edwards. [1863, Mar.-May - 12 Proc.
Ent. Soc. Phila. 2(1) : 14-15. Figs. 1-3 on plate 11 may have been
published later. "From Pike’s Peak; in the Society’s collection; 6
males, 1 female. The second female is from the collection of Mr.
George Newman and was taken among the Rocky Mountains . .
No dates given.
astraea, Colias W. H. Edwards. Feb. 1872. Trans. Amer. Ent. Soc. 4(?)
sign. 8: 61. "From a single male taken near the Yellowstone Lake
by the Hayden Expedition, io 1871”. No date given.
barbara, Colias Henry Edwards. 5 Feb. 1877’"^. Pac. Coast Lepid. (24) :
7-8 & 11. "Two females (Coll. Hy. Edw.), Santa Barbara, Baron
V. Osteo; Gilroy, J. Behrens”. [Santa Barbara & Gilroy are both in
California]. No dates given.
Christina, Colias W. H, Edwards. [12 Oct. 1863]’^. Proc. Ent. Soc.
Phila. 2(2): 79-80. 6 & 2 described. "Taken at the portage of
Slave River . . .”= No series data nor dates given.
chrysomelas, C{olias]. Henry Edwards. 5 Feb. 1877’"^ Pac. Coast Lepid.
(24): 8-9, 11. "Seven 6, ‘five 2. Napa County, Cal[ifornia]. (Coll.
Hy. Edw.)”. No dates given. Spelled chryomelas by H. Edwards,
Ibid., p. 9.
edwmdsii, Colias W. H. Edwards (Behr Ms. name). Jan. 1870. Trans.
Amer. Ent. Soc. 3(?) sign. 2: 11. "1 6,2 2, from the collection
of Dr. Behr, taken at Virginia City, [Nevada] at high elevation . .
No dates given. Spelled edwardsi by Barnes & Benjamin, 8 Dec.
1926, Bull. Sou. Calif. Acad. Sci. 25(3): 89, under 67a.
216
McHENRY
/. Res. Lcpid.
emilia, Colias W, H. Edwards. Jan. 1870. Trans. Amer. Ent. Soc 3(?)
sign. 2: 12. "From 1 6,1 2, taken in Oregon . . . Colleaion of
Dr. Behr”. No dates given.
harjofdii, C{olias\ Henry Edwards, 5 Feb. 1877'“^. Pac. Coast Lepid.
(24) ; 9 & 11. "From seven males, (Coll. Hy. Edw.) Contra Costa
Co. (Hy. Edw.). Havilah, Kern Co. (R. H. Stretch)”. [Both locali-
ties in California]. Spelled harfordi by Holland, 1931, Butt. Book,
rev. ed., p. 294, no. 5.
hatui, Eurymus alexandra edwardsi Barnes & Benjamin. 8 Dec. 1926.
Bull. Sou. Calif. Acad. Sci. 25(3): 89, under no. 67a. "Type
locality; Stockton, Utah. Number and sexes of types: Holotype $,
VII-30-16; 1 $ Paratype, VII-5-3.”
krauthii, Colias christina Klots. 24 Jan. 1935. American Mus. Nov. No.
767: 1-2. "Types. - Male holotype, allotype and thirty-one male and
seven female oaratypes, from twelve miles west of Custer, Black Hills,
South Dakota, June 29, 1933; four male paratypes, vicinity of Custer,
South Dakota, June 27, 1933, and five male and six female para-
types, Black Idills, South Dakota, July 1, 1931 . . "Holotype,
allotype, and twelve male and five female paratypes are in the
American Museum of Natural History; the remainder are in the
author’s colleaion.”
lambillioni^ Colias elis Dufrane. 28 Feb. 1947. Bull. & Ann. Soc. Ent.
Belgique 83(1-2); 70. 2 described. "... Canada, sans nom de
localite precise . . No date given.
martini, Eurymus harfordii Gunder. 10 Sept. 1931. Bull. Sou. Calif.
Acad. Sci. 30(2) : 45. "Holotype 2, . . . South side of Arrowhead
Lake, San Bernardino Co., Calif. Sept. 2, 1931. Type in Author’s
coll.”
occidentalis, Colias Scudder. Sept. 1862. Proc. Boston Soc. Nat. Hist.
9(?) sign. 7: 109-111; unnumb. fig. (p. 107). "2 6,3 2. Gulf of
Georgia (A. Agassiz); Fort Simpson, British America. (W. H.
Edwards,)”. No dates given.
pallida, C\plias\ alexmidra Cockerell. May 1889. Entomologist 22(311) :
128, under no. 2. Cites: "... (W. H. Edwards, Can. Ent., 1887,
229) . . Edwards speaks of two albino females sent by Bruce
[from Colorado?]. No dates given.
pallida, C[olias\ christina Cockerell. May 1889. Entomologist 22(311):
128, under no. 2. Cites: "... (H. H, Lyman, Can. Ent., 1884, 6)
. . Lyman speaks of two albino specimens of female. No dates
given.
palidissima, Colias eurytheme alberta Bowman. 9 Mar. 1942. Can. Ent.
74(2): 25. "Holotype - 2, Fort Vermilion, Al[ber]ta,, July 15,
1925, in the author’s collection. Paratypes - 2 2, same locality,
June 9 and 15, 1925, in the Canadian National Collection and the
colleaion of the author”.
shastae, Eurymus occidentalis chrysomelas Barnes & Benjamin. 8 Dec,
1926. Bull. Sou. Calif. Acad. Sci. 25(3): 88, under no. 63a.
"Albinic 2 of chrysomelas . . .”. "Type locality: Shasta Retreat,
Siskiyou Co., Calif. Numbers and sexes of types: Holotype 6 [in
error], 1-7 July”.
weaverae, Eurymus harfordii Gunder. 8 May 1924. Ent. News 35(5):
156-157; plate 2, fig. G. "Allotype [Holotype] 2, (Author’s Coll.)
Warner Sorings, San Diego County, California, July 3, 1949”.
9. COLIAS PALAENO ( LINNAEUS ) .
chippewa, Colias W. H. Edwards. [Aug. 1870]^'^. Syn. N. Amer. Butt,
(pp. 7-14) : 85 no. 15. A new name for his own Colias helena.
"Hab. - Great Slave Lake” [differs from that given for helena\ See
his helena for data.
(3):20^-22l , 196}
NAMES — Pt. 2
217
helena, Colias W. H. Edwards. [12 Oct. 1863]^^. Proc. Ent. Soc. Phila,
2(2) : 80. d & $ described. ''From Mackenzie’s River, . . No
series data oor dates given. A homonym of Colias helena Herrich-
Schafifer, 1844’'’. Syst. Bearbeit. Schmett. Europa 1(7): plate 45,
figs, 206-207 (I have not seen this reference of Herrich-Schaffer) .
kohlsaati, Eurymus chippewa Gunder. 10 Sept. 1931. Bull. Sou. Calif.
Acad. Sci. 30(2) : 45. "Holotype . Mt. McKinley Nat. Park,
Alaska. July 14, 1930”. "Type in Author’s coll.”
lapponica, Colias palaeno Staudinger. End Jan. 1871. In Staudinger
& Wocke. Catal. Euron. Faun., p. 5, no. 58a. $ described. Cites
figs. 403-4 ($, werdandi) on plate 83 in Herrich-Schaffer. 1848’'",
Syst. Bearbeit. Schmett. Europa 1(36). "Lap. Ross. s.”. No series
data nor dates given.
palaeno, Papilio [Heliconius] Linnaeus. 1761 [after 28 July]’®. Fauna
Svecica. Ed. Alt., p. 272, no. 1014. "Habitat in Pteride rarissime
Upsaliae, frequentior in Finlandia”. No sex data, series data nor dates
given.
philomene, [Papilio'] Hubner. [1805]’“^. Samm. Europ. Schmett. (Papili-
ones); no text; plate 117, figs. 602-603. Figs. 740-741 on plate 147
were ptiblished later.
werdandi. C[olias\ Herrich-Schaffer (Schdnherr Ms. name). 1844’^. Syst.
Bearbeit. Schmett. Europa 1(3): plate 8, figs. 41-42; 1(7): 102
(published 1844); plate 83, figs. 403-404 (published 1848). No
series data, no locality nor dates given. Homonym of Colias
werdandi Zetterstedt, Insecta Lapponica, page 908, no. 2, [1839]^°.
10. COLIAS PELIDNE BOISDUVAL & LECONTE.
interior, Colias Scudder. Sept. 1862. Proc. Boston Soc. Nat. Hist. 9(?)
sign. 7: 108-109; 1 unnumb, text fig. (p. 107). "5 $, I 2.
Northern shore of Lake Superior (Prof. Agassiz); mouth Sasketch-
ewan River, British America (S. H. Scudder.)”. No dates given.
isni, Eurymus pelidne minisni Barnes & Benjamin. 8 Dec. 1926. Bull.
Sou. Calif. Acad. Sci. 25(3): 89, under no. 69c. "Type locality:
Laggan, Al[ber]ta. Number and sexes of types: Holotype $, 16-23
Aug.”
lahradorensis, Colias Scudder. Sept. 1862. Proc. Boston Soc. Nat. Hist.
9(?) sign. 7: 107-108; unnumb. fig. (p. 107). "8 d, 5 2.
Caribou Island, Strait of Belle Isle, Labrador. (A. S. Packard, Jr.)”.
No dates given.*
laurentina, Eurymus philodice Scudder. Mar. 1876. Proc. Boston Soc.
Nat, Hist. 18(2): sign. 12: 189-190. "... Cape Breton Island”
[from article title]. "Thirty-nine specimens were collected, of which
ten were gynandromorphic females, eight pallid females, and the
rest males”. No dates given.
minisni, Colias elis Bean. Apr. 1895. Psyche, Jour. Ent. 7(228): 228.
Intended to withdraw his Ms. name for what he considered the
albinic female of elis Strecker.
mira, Colias pelidne Verity. 1905-1911. Rhopal. Palaearctica; p. 347;
p. xxxiv; plate LXVIII, fig. 49; ol. LXVIII exolanation page, fig. 49.
"49 ... $ ... (Ravea, Labrador) [e coll. Elwes]”. No date given.
moeschleri, Colias pelidne Grum-Grshimailo. 30 July 1894. Horae Soc.
Ent. Rossicae 27 (3-4) : 379, no. 1. 1 2 described. "Specimen
unum e museo . . . Moeschleri, alterum, in Labrador anno
1889 collectum, e museo lepidopterologi gemanici Dom. M. Wiskotti
acceptum.”
nepi, Eurymus interior Barnes & Benjamin. 8 Dec. 1926. Bull. Sou.
Calif. Acad. Sci. 25(3) : 89, under no. 65. "Type localitv: Nepigon,
Ontario. Number and sexes of types: Holotype 2, and 12 Para-
type both 8-15 July.”
218
McHENRY
/. Res. Lcpicl.
fieri, Eurymus pelidne skinneri Barnes & Benjamin. 8 Dec. 1926. Bull.
Sou. Calif. Acad. Sci. 25(3): 89, under no. 69b. "Type locality:
Yellowstone Park, Wyo[ming]. Number and sexes of types: Holotype
$ 8-15 July; 2 $ Paratypes, 8-15 July and no date; all being
original type females of skinneri Barnes.”
pelidne, C[olias]. Boisduval & LeConte. 1829. Hist. Gen. Icon. Lepid.
Chen. I’Amer. Sept. 1(7): plate 21, figs. 4-5 (2). Pp. 66-67
published in livrason 8. 6 & 2 described. "Elle habite le Greenland
et rislande. M. Sommer, observateur tres-distingue, me I’a aussi
envoyee comme venant du Labrador”. No series data nor dates given.
skinneri, Colias pelidne Barnes. 1 Feb. 1897. Can. Ent. 29(2): 41-42.
"Described from 15 males and 7 females - three of which are yellow,
three white, and one intermediate — taken in Yellowstone National
Park, and at Arangie, Idaho, in July”.
solivaga, Colias W. H. Edwards (Henry Edwards Ms. name). 1877 [Feb. -
16 Apr.]^’. Trans. Amer. Ent. Soc. 6(?) signs. 1-6: 17, under no.
68. Gives a Henry Edwards Ms. name as a synonym of interior
Scudder.
11. COLIAS PHILODICE GODART.
alba, Colias philodice Strecker. 1878 [Sept. - Oct.j'^. Butt. & Moths N.
Amer., Compl. Syn. Catal., p. 82, no. 58a. Cites 2 figures in W. H.
Edwards, Butt. N. Amer. 2 : plate Colias II, fig. 6 and plate III, figs.
5-6, 1876. No locality, series data nor dates given here.
alba, Colias philodice plicaduta F. Chermock. 28 Apr. 1927. Bull.
Brooklyn Ent. Soc. 22(2): 119. "Holotype 2. July 30, 1924; 2
paratypes August 17, 1925; 1 paratype August 16, 1925. N. S.
Pittsburgh, P[ennsylvanila.”
albida, Eurymus philodice plicaduta F. Chermock. 15 Nov. 1928. Bull.
Brooklyn Ent. Soc. 23(4): 173. A new name for his own Colias
philodice plicaduta alba which he considered a homonym of Colias
philodice alba Strecker, 1878. See his alba for data.
albinic, Colias philodice W. H. Edwards. [1884, after 12 Sept. - before
18 Feb. 1885]^^. Trans. Amer. Ent. Soc. 11(3-4): 264, under no.
68. Cites his own Butt. N. Amer. 2; plate Colias II, 1876. No
locality, series data nor dates given here.
anthyale, Zerene Hiibner. 1823 [1 Jan. - 20 Apr.]^^. Ziitr. Samm. Exot
Schmett. 2nd. 100: 21, no. 154. Figs. 307-308 on plate [54] were
published earlier but without names. $ described. "Aus Pennsyl-
vanien”. No series data nor dates given.
autumnalis, Colias eurytheme eriphyle Cockerell. Mar. 1888 [after 9th]°.
West Amer. Scientist 4(35): 41-43 (in pt.). "On November 9,
1887, I caught a female specimen of the autumn brood of eriphyle
in this locality” [(West Cliff, Custer Co., Colorado)].
ehrmanni, Colias philodice plicaduta F. Chermock. 28 Apr. 1927. Bull.
Brooklyn Ent. Soc. 22(2): 118. "Holotype 2. August 3, 1921.
N. S. Pittsburgh, P[ennsylvani]a.”
eriphyle, Colias W. H. Edwards. Apr. 1876. Trans. Amer. Ent. Soc.
5(?) sign. 26: 202-204. "Some thirty individuals of this species
and of both sexes, were taken at Lake Labache, in British Columbia,
. . . , and the whole series came into my possession”. No dates given.
europome, E\apili6\. D[anaus'). C{andidus\ Haworth. 1803 [in or after
July]’®. Lepid. Britannica (1): 13, no. 12. "Habitat in Anglia,
rarissimo. D. Francillon et D. Swainson”. "Descriptio (Ex exemplario
Germanico)”. No date given. A homonym of europome Esper
[1777]13, Schmett. 1(1): plate 42, figs. 1-2 (I have not seen
this reference of Esper) .
hagenii, Colias W. H. Edwards. [1884, 19 Tan. - 20 Feb.]^"^. Papilio
3(7-10): 160-161, no. 11; 3(7-10): 163-164. "From 20 6 24 2
from various localities, from So. Colorado to Montana and Dacotah
l(}):209-321, 1963
NAMES ™Pt. 2
219
(Bismarck)”. No specific dates given. Spelled hageni by Holland,
1931, Butt. Book, rev. ed., p. 297, under 00. 17.
hybrida^ Colias philodice Strecker, 1878 [Sept. - Oct.]"^. Butt. & Moths
N. Amer., CompL Syn. CataL, p. 82, 00. 58d. 8 cited in W. H.
Edwards, 1876, Butt, N. Amer. 2: plate Coiias III. "Illinois,
Georgia”. No date given here.
invetsata, Colias philodice Nakahara. 27 Jan. 1926. Bull. Brooklyn Ent.
Soc. 20(5): 223. "Holotype: $, Midvale, Allegheny Co., P[enn-
sylvanija., August 20, 1893 . . • Type in the Barnes Collection.”
kootenai, Colias Cockle. 4 June 191(). Can. Ent. 42(6): 203-204.
"Kalso and in Northern British Columbia” [from article title].
"... May 17 to 20, and fall brood to Oct. 9”. No sex data nor
series data given.
laurae, Eurymm eurytheme eriphyle F. Chermock. 9 Mar. 1929. Bull.
Brooklyn Ent. Soc. 24(1) : 21. 9 described. "Holotype, VIII-1-
1924: Edmondton, Alberta, Canada.”
lutetincta, Colias philodice Wolcott. 11 Apr. 1893. Can. Ent. 25(4):
104, under no. 50. "I first met this form ... in August, 1885, at
Batavia, Ill'[inoi]s . . . B.ut have since taken it at different times at
Grand Rapids, [Michigan] . . . and have collected in all about a
do2e,o specimens, one of them a female”.
maria, C{olias\ philodice W. H. Edwards (Grote Ms. name). 13 July
1885. Papilio 4(9-10): 171. Inadvertently describes a cyanide
stained specimen.
melanic, Colias philodice W. H. Edv/ards. [1884, after 12 Sept. - before 18
Feb. 1885]^^. Trans. Amer. Ent. Soc. 11(3-4): 264, under no. 68.
Cites [$] in his Butt. N. Amer. 2: plate Colias III, 1876. No locality,
series data nor dates given here.
minor, Colias philodice plicaduta F. Chermock. 28 Feb. 1927. Bull.
Brooklyn Ent. Soc. 22(2): 118. "Holotype $. June 28, 1925.
N. S. Pittsburgh, P[ennsylvanPa.”
miscidke, E[urymm]. p\hilodice\ Scudder. 1 June 1889. Butt. East. U. S.
& Canada 2(8): 1115. "Mr. F. G. Sanborn has shown me a very
interesting suffused female of this species taken by Mr. John Osgood
in Lynn, Mass[achusetts]., in August, 1863’h
nig, Colias philodice Strecker. 1878 [Sept. - Oct.]"^. Butt. & Moths N.
Amer., CompL Syn. CataL, p. 82, no. 58b. S from Montreal, Canada
described; also cites W. H. Edwards, 1876, Butt. N. Amer. 2: plate
Colias III, figs. 8-9 (copied from Glover’s unpublished plate). No
dates given.
nigficosta, Eurymus eurytheme eriphyle F. Chermock. 9 Mar. 1929. Bull.
Brooklyn Ent. Soc. 24(1): 21. " . . . occurs in the female sex
only”. "Holotype, VIII-1-1924; Topoparatype, VIII- 1-1 924; Ed-
mondton, Alberta, Canada”.
nigridice, Eurymus phil[odice]. Scudder. 1 June 1889. Butt. East. U. S.
& Canada 2(8): 1114. d & $ described. Cites W. H. Edwards,
1876, Butt. N. Amer. 2: plate Colias III, figs. 8-9 (S ). Cites fig. 5
on Glover’s unpublished plate I. Also cites other specimens which
were not in his possession (one captured 29 July 1883).
nigrina, Colias philodice Strecker. 1900 [Mar. 9th - 13th]^. Lepid.,
Rhopal. & Heter., Indig. & Exot. SuppL No. 3: 19. Applies the
name to what he formerly called nig. Here he cites 3 specimens:
"... Bethlehem, P[ennsylvanPa. . . . Montreal, Cao[ada] . . . Orillia,
Can.”. No dates given.
nigrofasciam, Colias philodice Reiff. 15 Jan.- 1917. The Lepidopterist,
Off. Bull. Bost. Ent. Club. 1(3): 22-23; plate 3, 1 unnumb. fig.
2 described. "... was caught . . . during August, 1916 in Massa-
chusetts, and it is now in the collection of the author”.
220
McHENRY
/. Res. Lcpid.
notatus, Papilio Megerle. 28 Nov. 1803. Catalogus Insectorum Qua
Viennae Austriae 28. Novembris 1803 Auctionis Lege Distrahuntur,
p. ?, no. 436. "M. ex. Georg[ia] . . . 6’’. No dates given. I have
not seen this reference, all data is from Clark & Clark.
notatus, Papilio Clark & Clark (Megerle Ms. name). 30 Apr. 1941.
■Proc. Ent. Soc. Washington 43(4) : 81, no. 436; & page 83, no. 1.
"M. ex Georg[ial. . . 6”. No dates given. Reprint Megerle’s
description.
pallidice, E{urymus\ p[hilodice\. Scudder. 1 June 1889. Butt. East. U. S.
&c Canada 2(8) : 1115; pi. 7, fig. 6; pi. 7 explanation page, fig. 6.
Page 1125 (published later). $ described. Cites W. H. Edwards,
1876, Butt. N. Amer. 2: plate Colias II, fig. 6 and plate Colias III,
figs. 5-6. Also cites fig. 11 on Glover’s unpublished plate 6. Data
is general.
philodice, Colias Godart. 1819. In Godart & Latreille in Latreille. Encycl.
Meth. 9(1) : 87, no. 35 (named in French); 9(1) : 100-101, no. 35.
[6] & $ described. "File se trouve dans I’Amerique septrentionale,
particulierement dans la Virginie”. No series data no dates given.
Spelled philodin by W. H. Edwards, 1872, In Hayden, Prelim. Rept.
U. S. Geol. Surv. Mont. & Port. Adj. Terr. 5th. Rept. Progress, p. 466.
plicaduta, Colias philodice Nakahara. 27 Jan. 1926. Bull. Brooklyn Ent.
Soc. 20(5) : 222-223. "Holotype: $, Ithaca, N. Y., July 30, 1923
. , . Paratopotype : $, July 16, 1924. Paratype: $, Lava, Sullivan
Co., N. Y., "June.” Holotype has been presented to the Barnes
Collection, Decatur, Illinois]., and paratype is in the same collection.
Paratopotype is retained in the collection of the writer”.
raritus, Eurymus philodice Gunder. 30 July 1928. Can. Ent. 60(7):
163; plate A, figs. 2, 2a. "Data: Holotype $ . . . Scranton,
P[ennsylvani]a. (bred . . .), July 8, 1927. In the Author’s coll.”
reducta, C\qlias\ philodice anthyale Dufrane. 28 Feb. 1947. Bull & Ann.
Soc. Ent. Belgique 83(1-2): 69. $ described. ”. . . Temple,
P[ennsylvani]a. . . ”. No series data nor dates given.
fothkei, Colias philodice Reiff. 15 Aug. 1917. The Lepidopterist, Off.
Bull. Bost. Ent. Club. 1(11): 84; plate 7, 1 unnumb. fig. "Col-
lected August 27, 1905 in the Susquehana Valley, P[ennsylvani]a.,
. . . Type 1 male in Mr. Rothke’s collection”. Mentions another
"specimen (no status).
s errata, Eurymus philodice F. Chermock. 9 Mar. 1929. Bull. Brooklyn
Ent. Soc. 24(1): 21. "... occurs in tlie male sex only . . .”.
"Holotype, VIII-8-1926; paratypes. No. 1, VIII-8-1926; Nos. 2, 3, 4,
VII-10-1926; No. 5, VIII-12-1926; Rossgrove, near Aspinwall,
P[ennsylvani]a.”
suffusa, C[olias]. philodice Cockerell. Mar. 1889. Entomologist 22(310):
55. Cites: ". . . Massachusetts, Maynard, Butt, of New Eng., pi. VII,
fig. 57c”. No date given.
virida, Colias philodice Strecker. 1878 [Sept. - Oa.]"^. Butt. & Moths N.
America, Compl. Syn. Cat., p. 82, no. 58c. $ described. "One
example . . Type locality: [near Montreal, Canada]? No date
given.
vitahunda, Colias chrysotheme Hovanitz. 30 July 1943. American Mus.
Nov. No. 1240: 2, 3-4. "Holotype Female and Alotype Male. —
McKinley National Park, Alaska, July 18 to August 9, 1930, . . .,
J. D. Gunder Collection, in the American Museum Natural History.
Paratypes. — Twelve males same data as above; eleven males same
locality but July 29, 1931, . . .; for males July 15-20, 1931, ... in
Los Angeles Museum; ten while female same data as holotype; four
white females July 20-30, 1931, ... in Los Angeles Museum; one
white female August 9, 1930, ... in Los Angeles Museum”.
l{})X20^-22l , 196}
NAMES — Pt. 2
221
12. COLIAS SCUDDERII REAKIRT.
flavotincta, Colias scudderi Cockerell. Apr. 1901. Psyche, Jour. Ent.
9(300) : 186. Cites W. H. Edwards, Butt. N. Amer. [1]: plate 10,
fig. 5 (?) (part of illustrated series of scudderii, 1872).
ruckesi, Colias scudderi Klots. 2 Nov. 1937. Jour. New York Ent.
Soc. 45(3-4): 324-326. "Holotype male, allotype female, nineteen
male and one female paratypes, from Windsor Creek Canyon, west
of Cowles, N. Mex., July 2, 1935, . . . Twenty-five male and nine
female paratypes from the same locality, July 4, 1936, . . . All were
taken in about middle Canadian Zone, at from 9000 to 9500 ft.
altitude, in grassy meadows surrounded by forest, along the Forest
Service trail about halfway between Cowles and the summit of
Santa Fe Baldy Peak. Holotype, allotype, six male and two female
paratypes deposited in the American Museum of Natural History;
four male paratypes deposited in the U. S. National Museum; four
male paratypes deposited in the Canadian National collection; the
remainder of the paratypes at present in the author’s collection.”
scudderii, Colias Reakirt. [12 June 1865]^^. Proc. Ent. Soc. Phila. 4
(pp. 213-339) : 217-218, no. 2. "Hab. — Rocky Mountains, Colo-
rado Territory. (In my collection, 8, and that of the Entomological
Society. $.)”. "This species was collected in August, 1864 . . .”.
Spelled scudderi by Holland, 1931, Butt. Book, rev. ed., p. 295,
no. 10.
FOOTNOTES
IVol. 6 title page date is qu*Iified by indirect date data (certain names in Fabricius’ article
were mentioned in the Allgemeine Literatur-Zeitung, Halle [Rnal 2(303): cols. 1177-1181, 19
Dec. 1807).
2Pt. 2 title page date is qualified by indirect date data by A. S. Corbet, 28 Feb. 1939, Jour. So.
Bibliop-. Nat. Hist. 1(7): 96.
^Hemming. 1937. The Generic Names of the Holarctic Butterflies 1: 139, see no. 376.
"^Work title page date is qualified by indirect date data on pages 39 & 48 in the Sept. & Oct.
numbers of Vol. 3 of the Bull. Brooklyn Ent. Soc.
5Dos Passos. 1962. Jour. Lepid. Soc. 16(1): 45-46.
^Sherborn & Woodward. 1901. Ann. & Mag. Nat. Hist. Ser. 7. 7: 137-140.
7No. 3 title page date is qualified by the preface date and a Ms. date of receipt (13 Mar. 1900)
written on a copy of No. 3 in the library of the Los Angeles County Museum.
®No. 35 title page date is qualified by the article signature date.
9The date Sept. 1869 (given for Pt. 4 as per footnote 16) is qualified by the receipt date or
Pt. 4 (see Trans. Amer. Ent. Soc. 2; XVIII, 1868-9).
’OTome 5 title page date is qualified by a meeting date (see p. 383).
^ IKirby. 1871. Syn. Catal. Diurn. Lepid., p. 494, no. 29a.
1 ^Article received by publisher 9 Mar. 1955. No. 1 containing the article was received at the
Allen Hancock Foundation Library (Univ. Sou. Calif.): 11 Oct. 1955.
l^No. 1 title page date: Mar., Apr. & May 1863. No. 2 receipt date (see Proc. Ent. Soc. Phila.,
2: 164).
14More actual publication date data is needed.
'5Proc. Ent. Soc. Phila. 2: 164 (No. 2 receipt date is given). No. 2: pp. 61-162.
1 ^Hemming. 1931. Proc. Ent. Soc. Lond. Ser. A. 6: 42-45. Pp. 7-14 of Synopsis were published
with Butt, N. Amer., pt. 6.
17Hemming. 1937. Hubner 1: 579-589.
1 ®Title page date is qualified by the preface date.
^ ^Hemming. 1937. Hubner l:146-3Z‘t.
20Sherborn, 1922. Index Animalium, 2nd. Ser. Vol. A-B: cxxxi.
21 Vol. 6 title page date is qualified by the date of signature 3 and the date of a letter of W. H.
Edwards to H. Edwards (Henry Edwards correspondence in the Amer. Mus. Nat. Hist, N. Y.)
dated 16 Apr. 1877 and stating he would mail a copy of the catalogue the next day.
22No. 2 (Vol. 11) of the Trans. Amer. Ent. Soc. was available 12 Sept. 1884 (see p. xxxvi,
vol. 11). The issue date of the separate was 18 Feb. 1885 (see page 2 of separate).
23Hemming. 1937. Hubner 1: 451-487.
24Nos, 7-10 (v. 3) were not published before 19 Jan. 1884 (see indirect date data on page 193).
No, 1 (v. 4) was published 20 Feb. 1884 (see p. 42, v. 4).
25pp, 221-330 (Mar. & Apr. 1865) receipt date (see Proc. Ent. Soc. Phila., v. 4, p. xii).
y
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Journal of Research on the Lepidoptera 1 (3) :223-235, 1963
1140 W. Orange Grove Ave., Arcadia, California, V.S.A.
© Copyright
A STANDARD METHOD FOR MOUNTING
WHOLE ADULT LEPIDOPTERA ON SLIDES
UTILIZING POLYSTYRENE PLASTIC
CHARLES L. HOGUE
Los Angeles County Museum, Los Angeles 1 , California
Comparative morphological studies of insects are greatly
facilitated if the material is prepared and mounted in a way that permits
its easy manipulation and rapid, yet detailed, examination. Mounting
specimens dry, on pins, has been the method of preference of Lepid-
opterists virtually exclusively, but it does not permit detailed examina-
tion of integumentary features that are of great taxonomic importance.
Attempts to interpret the relationships of higher categories in the
lepidoptera entirely on the basis of structure externally visible on such
specimens have, in many cases, led to errors and have confused the
taxonomy of many groups.
Although most workers are willing to utilize slide mounts for limited
parts of the body, especially the genitalia, they have not extended the
treatment to the whole animal and prefer to keep their material in
fluids, usually alcohol. While alcoholic specimens allow the greatest
amount of freedom of examination, considerable time and patience
is spent simply handling them — ^unstoppering vials, pouring alcohol,
restoppering vials, etc. As a result, a worker may hesitate to make the
innumerable comparisons and recomparisons requisite to phylogenetic
analysis. Furthermore, since the specimen must be examined with
relatively low power dissecting microscopes, many fine details, such as
microchaetae and other minute sense organs, surface textures, striae,
spicules, spiracular structure, etc., are not visible, or at least not clearly
observable. Among the other disadvantages of alcoholic specimens are*
that they do not keep stains well, are subject to evaporation and destruc-
tion from drying and suffer damage due to repeated handling.
Slide mounts, on the other hand, permit very detailed observation
and are very easily handled. Admittedly, they allow the specimen to
be viewed only from a limited number of aspects, but if prepared in
a careful manner, such as that described below, most of the integu-
mentary anatomy is available for inspection. Slides may be stacked as
high as 4 or 5 deep and the specimens readily compared by focusing
up and down with a dissecting microscope or detailed drawings rapidly
prepared for the same purpose with a microprojector. In addition, they
may be permanently stained, are subject to destruction only by breaking,
223
224
HOGUE
/. Rex. Lepid.
and may be handled indefinitely without wear. Even fairly large Lepid-
optera (e.g., smaller Catocala, Vanessa) may be placed on slides but I
do not recommend it for very large species (e.g., saturniids, sphingids).
The details of the method outlined below were worked out on
noctuid moths of medium size with relatively simple genitalia. Yet I
believe that most groups of Lepidoptera may be similarly treated,
although, of course, certain groups may require special handling be-
cause of extremely small size or peculiar modifications, especially in
the genitalia.
The general treatment below might be applied advantageously to
many insect orders (especially Neuroptera, Trichoptera, Diptera, etc.).
All stages of mosquitoes are beautifully accommodated and I have had
promising results with Collembola, a group notoriously difficult to
mount on slides. The latter usually shrivel badly in balsam and eventu-
ally deteriorate if placed in aqueous media, apart from being flattened
or otherwise distorted.
THE USE OF POLYSTYRENE PLASTIC
Whole mounts of most insects require fairly thick preparations.
Workers who have tried to make such preparations with the usual
natural (Canada balsam, Euparal), synthetic (Permount, Diaphane),
or mixed aqueous (PVA, Hoyer’s) media find that there are certain
difficulties: These media, when applied in large amounts take a very
long time to -harden, specimens invariably move in them from their
desired positions during the hardening period (even if the laborious
and time consuming "layering” or "build-up” technique is used), and
they change color or deteriorate after a time.
Polystyrene plastic' (hereafter referred to simply as "plastic”),
however, primarily because of its setting qualities, is the ideal medium
for holding dissected parts in their desired positions on slides. Because
it gels evenly and rapidly throughout, any specimen, regardless of shape
or center of gravity, may be orientated perfectly in any position. Further
advantages of plastic over the media mentioned above are its greater
transparency and resistance to discoloring, and its thorough hardness
and permanency when finally cured. In addition, it has the desired
qualities of being its own clearing agent and of slightly intensifying
certain stains. It has one disadvantage at present; it cannot be dissolved
gradually and without fragmenting after complete hardening by any
agent I know and therefore a specimen cannot be removed once
mounted. This is a serious drawback, however, only with scarce material
which might need to be remounted for study from a different aspect.
The methods described in this paper are intended primarily for morpho-
logical and phylogenetic studies where abundant material is available.
In any case, since slides are rarely remade unless very poorly prepared
'Known commercially by many trade names, this is the type used for embedding biological
specimens and other items by hobbyists.
(3j;225-2j5, 196}
PLASTIC METHOD
225
in the first place, the insolubility of plastic should not limit its use.
Preparations on broken slides may be soaked off in cellosolve and glued
onto new slides with a thin layer of plastic.
The steps involved in using plastic as a medium for thick micro-
scope slide mounts are as follows:
1. The specimen, after dissection and dehydration in up to 95% ethanol
(see directions below), is transferred to a mixture of 50% acetone and 50%
uncataly2ed plastic (because of the difference in density between these compon-
ents they do not mix readily- — -mixing may be accomplished by shaking them
together in a vial for a few seconds). Since the acetone evaporates rapidly
from the mixture, its container must be kept covered as much as possible. The
mixture clears and completely dehydrates the specimen; the specimen should
remain in it about fifteen minutes.
2. While the specimen is clearing, a slide is prepared to receive it in the
following manner: Uncatalyzed plastic is dropped onto the slide and spread
out over an area the size of the cover slip. Spreading is done with a glass rod
(3mm. diameter), which has been previously dipped into catalyst. The rod is
used to apply and thoroughly mix the catalyst into the plastic. The depth to
which the rod is dipped determines the amount of catalyst picked up. I find
that dipping it about 5mm. provides the right amount of catalyst to gel the
thickest layer of plastic that can. be spread over a 22mm. square area (size
of standard, square cover slip) without running over. Enough plastic should
be applied to just cover the thickest part of the specimen. If the specimen is
very thick and the viscosity of the plastic is not enough for it to stand level
with the specimen, more layers must be added.
3. The specimen is next transferred to the plastic on the slide and
generally oriented.
4. The slide is then placed on a warming plate, previously heated to
about 130°F. to accelerate the gelling of the plastic. While the preparation is
on the plate, it is viewed with the low power of a dissecting microscope and
the specimen appropriately oriented. The specimen is held in its desired posi-
tion with dissecting needles until the plastic gels enough to firmly anchor it.
At this temperature, gelling will begin rapidly and will be far enough advanced
to hold the specimen in about 7-10 minutes^. Impending gelation can be noted
by the cessation of movement in the plastic revealed by dust particles present
in it.
5. The slide is allowed to remain on the warming plate about 15 minutes
to ensure thorough gelation.
6. The preparation is then ready to receive a cover slip if the first layer
of plastic was sufficient to cover the specimen. If not, more layers must be
added in the same manner as the first.
7. Glass chips or other cover slip supports are unnecessary and should
not be used since they hinder the settling of the cover slip on the slightly
shrinking plastic, and may cause a broken preparation »
8. The cover slip is prepared by covering it with just enough catalyzed
plastic (applied with glass rod as above) to spread over its entire area when
inverted and placed on the specimen. When the cover slip is in place the
whole preparation is returned to the warming plate for gelling. The heat also
helps the plastic under the cover slip to spread. If too little plastic was applied,
additional amounts may be added at the side of the cover slip until all the
spaces are filled. Any excess plastic may be wiped off from around the edges
or, if gelling has occurred, trimmed with a blade.
9. Complete hardening and curing of the whole preparation can be
accomplished by leaving it overnight in an oven at about 100°F. or can be
allowed to take place slowly at room temeprature.
2This figure varies according to the formulation and age of the plastic. I find these times average
but they may be longer especially for formulations designed for long shelf life.
226
HOGUE
/. Res. Lepid.
STANDARD DISSECTION AND MOUNTING
Four slides are required to hold the various parts of the whole insect
(fig. 1): 1. The main part of the body including the two halves of
the head, thorax and abdomen are put on the first slide (the tegulae,
antennae and basal portions of the right wings being dissected away— -
see below) (fig. 1:1); 2. A second slide holds the legs, basal portions
of the right wings, antennae and tegulae (fig. 1:2); 3. The wings
occupy a third slide ( fig. 1:3); 4. The genitalia are put on a final
slide (be they male (fig. 4b) or female (fig. 4a).
These main parts are dissected and mounted as follows:
Preliminary general dissection
The following instructions apply to specimens mounted on pins. Specimens
in alcohol or otherwise preserved are excused from the impertinent steps.
1. The left pair of wings are broken off at their bases and the right pair
cut off transversely with scissors beyond the end of the frenulum. The wings,
with the data labels removed from the pin and a slide reference label, are set
aside in a cellophane envelope.
2. The body of the insect, still on its pin, is wetted in 75% ethanol
and then boiled in water for 5 minutes. This softens it and allows the pin to
be removed easily.
3. The body is next transferred to a solution of 10% potassium hydrox-
ide which is heated to near boiling in a small beaker or test tube suspended in
a water bath. The specimen is left in the solution until thoroughly macerated
( 15 minutes) .
4. After maceration, the body is transferred for dissection and cleaning
to distilled water in a watch glass, preferably one with a slightly convex
bottom.
5. Next the genitalia are removed. In the male this is done by severing
the membrane between abdominal segments VII and VIII (or further cephalad
to include special modifications on more proximal abdominal segments if
present). The female is treated likewise, but by severing the membrane between
segments VI and VII. The genitalia are rubbed clean of scales (I like to use
a pointed applicator stick for this; others may prefer a fine camel’s hair brush
or like instrument) and transferred to clean distilled water.
6. The legs are now removed (coxae and trochanters should remain
intact), rubbed clean of scales and transferred to clean water.
7. The body itself (head, thorax with bases of legs, and abdomen minus
genitalia) is finally bisected with a sharp, single-edged razor blade. Great care
must be taken at this step to insure as perfect a bisection as possible. The
specimen is held ventral side up in the watch glass and a mid-ventral, longi-
tudinal slit is made at the posterior end of the abdomen with scissors. The
razor blade is then brought down on the mid-ventral line guided by this slit,
the coxae and the two halves of the proboscis between which its edge is
placed. The blade is first used to compress the body gently and then pressed
firmly against the bottom of the watch glass and rocked to and fro to make
the final cut. The slightly convex bottom of the watch glass helps to make a
clean cut. After the halves of the body are separated, cleaned and rubbed free
of scales, they join the other parts in distilled water.
8. Dehydration in 75% and 95% ethanol follows (if no aqueous stain
is used— -see "STAINING”). The female genitalia and phallus of the male
genitalia go dirertly to 95% ethanol to harden their membranous parts. This
is done in a special manner as described below^
fj).- 223-23 5, 1^63
PLASTIC METHOD
227
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Fig. 1: Set of slides (I - 4) holding a whole specimen {4a example
a female; 4h a male) .
228
HOGUE
/. Res. Lepid.
(3):22}-2}5, 1^63
PLASTIC METHOD
229
Figs. 2-6: Close-up views of slides shown in fig. 1 to show detail of
arrangement of parts. Fig. 2: Main body, bisected. Fig. 3: Miscellaneous
parts. Fig. 4: Wings. Fig. 5: Female genitalia. Fig. 6: Male genitalfa.
230
HOGUE
/. Res. Lcpid.
9. The wings, previously set aside in a cellophane envelope, are bleached
for about 2 or 3 minutes (at least long enough for the veins to become well
defined) in 5% sodium hypochlorite (full strength household bleach — Purex,
Chlorox), washed in distilled water, and dehydrated by passage through 75%
and 95% ethanol.
If it is desired that the veins be stained a totally different procedure
is necessary:
After wetting in ethanol and soaking for 20-30 minutes in hydrogen
peroxide ( 3 % ) , the scales are carefully rubbed off with a blunt camel’s hair
brush. The denuded wings are then washed in distilled water and transferred
to the stain. A long destaining period must follow in order to decolorize the
membranes.
The dissected parts, now in 95% ethanol, are ready for fine, detailed
dissection and cleaning (and staining with non-aqueous stains if desired — see
"STAINING”) before mounting in plastic on slides. I find that these two
jobs are best done in the 95% ethanol; since the specimen has become fairly
brittle, sclerites are easily separated and remaining unwanted scales easily
flaked off.
Final dissection and mounting
The different parts are treated in various ways. Reference to the
figures will clarify the descriptions (including terminology) of the fine
points of dissection and orientation on the slide.
Slide 1: Main body (figs. 1:1;2)
The two halves of the main body of the specimen require only
minor alteration before being transferred to the clearing mixture. The
tegulae and antennae are removed and the bases of the right pair of
wings are carefully dissected away with the axillary sclerites. These
parts are mounted with the legs (see "Slide 2”). If the specimen is a
noctuoid moth, the ventral portion of the second phragma is also cut
away with fine scissors to allow an unobstructed inner view of the
tympanal area. Similar special treatment may be necessary with other
taxa.
The halves are mounted on the slide, one with its outer surface up
and the other (the one with the cut phragma if noctuoid) with its
inner surface up.
Slide 2: Legs, etc. (figs. 1:2;3)
The legs, bases of wings (with axillary sclerites), tegulae and
antennae are all mounted together on the same slide since they are of
about the same thickness and do not orient well if left attached to the
main body. Usually these parts require no further dissection although
sometimes one may desire to separate the components of the pretarsus
or the antennae. Their orientation is best explained by the figures.
Slides: Wings (figs. 1:3;4)
No special treatment is needed for the wings. After dehydration
they are simply mounted flat in the plastic ( which also clears them ) .
If too large to fit on a standard microscope slide, they may be accom-
modated by a large sized .slide (2"x3^')- if very large, they may have
to be cut up into sections.
if3j-"225-2j5, 1963
PLASTIC METHOD
231
Slide 4: Genitalia — female (figs. l:4a;5;7)
The following procedure is considerably modified from that usually
employed by lepidopterists. The conventional procedure is to leave
abdominal segment VII attached and to mount the entire undissected
complex ventral side up or laterally on the slide. This, however, has
the following undesirable features: (1) segment VII, still attached,
obscures detailed viewing of the structures beneath, (2) if mounted
with the ventral side up the venter of segment VIII is plainly visible
but the pleura, which also may possess important characters, are not.
Furthermore, only the ventral edges of the ovipositor lobes are in plain
view; their profile, patterns of sclerotization and chaetotaxy are impos-
sible to see. If mounted laterally, the reverse viewing difficulties are
present. (3) The whole complex is cylindrical in general shape and
rolls easily, making it hard to position in a standard way.
By mounting the female genitalia as described below, these diffi-
culties are surmounted. The steps involved are as follows (fig. 7 shows
them partly completed) :
1. The genitalia have been freed from the remainder of the abdomen
and cleaned of most of their scales (see above, "Preliminary general dissec-
tion”)-
2. Segment VII is removed by splitting it laterally and tearing it off. The
tear should follow the line of the membrane between it and the following
segment. Care must be taken not to damage the sclerites around the ostium
bursae.
Fig. 7 : Oblique dorsal view of female genitalia showing principal
structures in state of partial dissection (diagrammatic).
232
HOGUE
/. Res. Lepid.
3. Next, the genitalia proper are slit mid-dorsally, a cut being made
first with scissors through segment VIII and continued by tearing caudad to the
tip between the ovipositor lobes.
4. The genitalia are then opened up, the oviduct and rectum are pulled
up and cut free, and' the whole complex cleaned.
5. The bursa copulatrix is next inflated with water in much the same
manner as the vesica of the male phallus (see below). The cannula is inserted
into the ostium bursae and passed down the ductus bursae. The stream of
water from the syringe cleans the corpus bursae internally and expands it
into its full shape. Sometimes a tiny hole is needed to allow the escape of the
fluid and debris. It is usually not necessary to go through a hardening process
with the bursa copulatrix as is the case with the male phallus vesica since it
generally holds its shape better than the latter,
6. Staining with aqueous stains follows if desired.
7. Preparatory to dehydration in 95% ethanol the genitalia are opened
again and placed flat (inner surface down) in a stender. A small piece of
glass (broken microscope slide) is placed over the opened portion (not over
corpus bursae! ) to hold it flat while the water is withdrawn from the vessel
and replaced with 95% ethanol. The same may be done for segment VII,
previously removed. It may be necessary, if not already done, to punch a small
hole in the corpus bursae before adding the ethanol if the ductus bursae is
so narrow that it would prevent the free exchange of liquids. Otherwise, the
diffusion gradient — ethanol diffusing out faster than water diffusing in — ^may
cause the collapse of the bursa.
8. Final cleaning may be carried out after dehydration. The preparation
is then ready for clearing and mounting.
Slide 4: Genitalia — male (figs. 1:4b; 6; 8-10)
Lepidopterists disagree on dissection procedures and orientation
of the male genitalia on slides. The majority prefer to leave all the
elements intact and mount the complex ventrally with the valves spread
open as widely as possible, the least troublesome method. Others also
do not separate the parts but mount the whole complex laterally. Still
others dissect various parts away, often only the phallus whose vesica
may or may not be inflated and the main orientation is either ventral
or lateral.
These variations may be dictated by either the whims of the investi-
gator or the nature of the material, i.e., the positions of the characters
of greatest significance. Too often it is the former. Obviously, the only
scientific and rational approach to the problem is to dissect and mount
many specimens in many ways, even keeping some unmounted in
fluids. Yet when time is limited, and many comparisons are to be made,
a single, standard method must be chosen. For this, I prefer to mount
the main structure of the genitalia laterally, with one valve and other
elements such as the juxta, anellus, etc. removed and mounted separately.
Furthermore, I always inflate the vesica of the phallus; though this is a
time consuming task, the results are well worth it.
Even apart from the conditions just mentioned, I believe that this
procedure is superior to the undissected ventral mount for morpholo-
gical study since there are generally better, undistorted viewing aspects
for the majority of characters and additional characters are revealed.
Specifically, (1) the uncus is natural in shape and position instead of
i(}):22}-2}5, 196}
PLASTIC METHOD
233
being folded or angled if long, or viewed on end if short. Its entire
profile is observable in plane view; (2) the tegumen is untwisted
( otherwise usually resulting from the disturbed uncus ) ; ( 3 ) the overall
shape, lobes and spines etc. of the vesica are fully exposed, not com-
pressed into the body of the phallus; (4) the anellus is not obscured
by the juxta (and phallus too if left intact^; (5) the valves (at least
the one dissected away and usually also the one intact) lie completely
flat so that their median structures are not distorted by foreshortening
as is usually the case in a ventral mount.
The steps followed in dissecting the male genitalia are as follows:
1. The genitalia have been freed from the remainder of the abdomen
and cleaned of most of their scales (see above, ''Preliminary general dissection”).
1 2. The phallus is removed by grasping it at the tip, i.e., posterior end
protruding between juxta and anellus (fig. 8), and pulling gently to the rear
(fig. 9). If the phallus does pot so protrude, the forceps may need to be
inserted a short distance into the canal inclosing it to gain a hold. If the
phallus is withdrawn properly and carefully, the manica will turn wrong side
out and the ductus ejaculatorius will pull out through the hole in it (fig. 9).
If staining with aqueous stains is desired at this point, the main portion
of the genitalia should be set aside in distilled water. Otherwise, they may go
directly into 95% ethanol.
3. The ductus ejaculatorius should now be snipped off with a pair of
fine scissors a short distance from the aedeagus,
4. Next comes the preliminary inflation of the vesica. This is accom-
plished by first teasing as much as possible of the vesica out of the aedeagus
with fine forceps and then inflating it with a hypodermic syringe filled with
distilled water. I find that a 2cc. syringe tipped with a glass cannula (in
preference to a regular hypodermic needle which does not permit the variety
of sizes needed) best for this work. The cannula is inserted into the ductus
ejaculatorius (fig. 10) and pressure applied until the vesica completely inflates
and everts. It is usually necessary to crimp the ductus with forceps against the
cannula at the point of insertion to prevent leakage and the phallus from
coming free under the hydraulic pressure.
5. The vesica is next hardened to preserve its fully inflated shape (fig.
10). This step follows staining if aqueous stains are used. Several steps are
necessary in this process:
a. The phallus is placed in clean distilled water in a watch glass.
b. A short length ( 1 cm. ) of silk surgical suture ( cardiovascular
5-0 or 6-0) is tied into a loose, simple knot and also put into the watch glass.
c. The hypodermic syringe is filled with 95% ethanol.
d. The cannula is inserted as above. This may be difficult since the
alcohol rapidly diffuses out of the tip setting up a strong current which
constantly carries the phallus away,
e. After the cannula is properly inserted the phallus is tied to it
with the suturing thread (around the center of the aedeagus) by slipping the
loose knot over it and drawing it tight with forceps (fig. 10). Some kind of
clamp or modelling clay is useful at this point to hold the syringe while both
hands are being used to tie the knot,
f. The water in the watch glass is now withdrawn and quickly
replaced with 95% ethanol.
g. Pressure is exerted with the syringe and maintained for a short
while (ab^out 3 minutes usually suffices) until the vesica is thoroughly
dehydrated and hardened. Leakage of the ethanol from the end of the vesica
is prevented by clamping it with fine forceps (fig. 10),
h. The phallus is slipped off the cannula and the suturing thread
untied and removed.
i. The phallus joins the rest of the genitalia in 95% ethanol.
234
HOGUE
/. Res. Lepid.
8
Figs. 8-9: Ventral views of male genitalia showing principal parts and
procedure for removal of phallus. Insert showing relationship^ of components
of phallus complex in lateral, sectional view. (All figures diagrammatic).
(3j.-223"2j5, 1963
PLASTIC METHOD
235
10
Fig. 10: Phallus rf male genitalia properly prepared for inflation and
hardening of ¥esica (diagrammatic).
6. Further dissection, if desired (at least segment VII 'should be
removed) and final cleaning may now be carried out in preparation of clearing
and mounting.
STAINING
Only a few general remarks regarding stains will be made here.
Each worker has personal preferences and many stains are in popular
use, among them add fuchsin, mercurochrome, and safranin. I have
always found the first the most satisfactory. Add fuchsin is taken up in
the greatest amounts by the sderotized parts of the integument leaving
membranous parts dear. Thus sderites are clearly defined and easily
observed and drawn. Tiny sclerotic processes in membranes also show
up very distinctly. I have not found the other stains mentioned to act
in this manner.
Add fuchsin is an aqueous stain, i.e., it is soluble in water and must
be applied to a fully hydrated specimen. I usually apply it after pre-
liminary dissection using a 5% solution diluted 1 drop to 10‘ ml.
Overnight is usually long enough to leave the parts in the stain. If not
enough stain is taken up in this time more is added until the specimen
is deeply tinted. Staining is always followed by destaining in distilled
water to clear the membranes and obtain the desired differentiation.
Non-aqueous stains may be applied to the specimens after they
have been dehydrated in 95 % or absolute ethanol.
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BIO METAL ASSOCIATES
Box 61
Santa Monica, California
Volume 1
Number 3
March, 1963
THE J0UIRNJAL ©F KESIA.^CH
©NJ THE LEFIJ©©FTE^A\
IN THIS ISSUE
Selection of allyl isothiocyanate by larvae of Pieris rapae and
the inheritance of this trait
William Hovanitz and Vincent C. S. Chang 169
Biology of the Ceanothus stem-gall moth, Periploca ceanothiella
(Cosens), with consideration of its control. J. Alex Munro 183
Larval food-plant records for six western Papilios
John F. Emmel and Thomas C. Emmel 191
Colias philodice in Chiapas^ Mexico . . Thomas C. Emmel 194
Notes on the early stages of two California geometrids
John Adams Comstock 195
Geographical distribution and variation of the genus Argynnis
III. Argynnis diana William Hovanitz 201
The generic, specific and lower category names of the nearctic
butterflies. Part 2 — The Genus Colias . Paddy McHenry 209
A standard method for mounting whole adult lepidoptera on
slides utilizing polystyrene plastic . . Charles L. Hogue 223
SPECIAL NOTICE
The Journal is starting a series of short illustrated biographical sketches
of past and present lepidopterists. In order to be wholly democratic and with
no aim in mind of "honoring” anyone, all lepidopterists will be included. The
only criterion will be a continued interest in the Lepidoptera, as shown by
publication, collections, etc. The editor would appreciate receiving black and
white prints of lepidopterists, together with a brief biographical sketch. This
applies to lepidopterists of all countries. Authors sending manuscripts might
send this material along with their articles.
Volume 1
Number 4
Mny,
THE JOURNAL
OF RESEARCH
ON THE LEFIJPORTERA
a quarterly published at
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
edited by: WILLIAM HOVANITZ
THE PURPOSE OF THE JOURNAL is to combine in one source
the work in this field for the aid of students of this group of insects
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AUTHORS ARE REQUESTED to refer to the journal as an
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THE JOURNAL is not a suitable place for continued changes
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Journal of Research on the Lepidoptera 1 (4) ;237-244, 1963
1140 IT. Orange Grove Ave,, Arcadia, California, U.S.A.
© C&py fight
TECHNIQUES IN THE STUDY OF
POPULATION STRUCTURE IN
PHILOTES SONORENSIS
RUDOLPH H. T. MATTONI and MARVIN S. B. SEIGER
Life Sciences Department, North American Aviation, Downey, Calif, and
Department of Biological Sciences, Purdue University, Lafayette, Ind.
The handling of living material has always been an im-
portant factor in the design of biological experiments. This factor is
especially critical in studying population structure under natural
conditions. One of the most accurate techniques for determining
parameters of population structure is that of capturing, marking, re-
leasing and subsequent recapturing of individuals (Ford 1951). Such
experiments would be biased if a behavioral response is elicited or
differential viability imposed as a result of the technique employed.
We have developed a handling technique in sampling populations
of the small Lycaenid butterfly, Philotes sonorensis, over a period of
three collecting years. This routine appears to have little or no effect
on the subsequent behavior of the butterfly. The purpose of this
report is to describe and evaluate the technique.
Our objectives were to describe the distribution, numbers, and
movements of adult individuals of P. sonorensis within a small circum-
scribed area in the Fish Canyon portion of the San Gabriel Canyon
Wash near Los Angeles. These individuals were classified as to 8 male
and 5 female spot pattern phenotypes. (Figure 1). Six stations, each
80 meters in diameter, were set up and sampled in 1955 and 1956.
These were separated by distances ranging from 96 to 433 meters from
the center of one area to the perimeter of another. The stations were
destroyed by trenching operations in 1961 because of water require-
ments. New experimental sites were established in 1963 in other
areas of the wash.
Sampling was done as weather permitted during the flight period
in March. In 1955 a total of 809 specimens were captured 1126 times
during 9 collecting days over a 21 day period. In 1956 there were
972 specimens captured 1226 times for 11 collecting days over a 29
day period. Sampling was done between 9 A.M. and 3 P.M., one
hour being allotted to each station during each day. The order in which
the stations were collected on consecutive days was ramdomized in
order to minimize possible differential effects correlated with time
237
238
PHILOTES SONORENSIS
/. Ref. Lepid.
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AiATTONI AND SEIGER
239
Fig. 2. (below) Photograph showing equipment described in the text.
RHTM on right holding a specimen about to be placed in the recovery
chamber on the portable desk. Vials with specimens can be seen in the com-
partment and the CO2 tank behind. MSRS on left recording data.
Fig, 3. (above) Anaesthesis Tube.
240
PHILOTES SONORENSIS
/. Res. Lepid.
of day. Each station was sampled for 30 minutes. This allowed 10
minutes for moving between stations and 20 minutes for handling.
When a butterfly was caught it was removed from the net with a
T’ X 4” shell vial and the vial was plugged with a cotton stopper. We
avoided touching the specimens with anything other than the net
and the vial. After the collecting period the vials of butterflies were
assembled in the center of the collecting area. In all cases one of us
(RHTM) classified and marked the specimens while the other
(MSBS) recorded the data. This process took approximately 20 minutes
per station. The equipment used is shown in Figure 2. This included
a portable desk, carbon dioxide tank, and release carton. The com-
partment on the desk served to store vials in the shade. Each specimen
was anesthetized in its vial by a 10 second exposure to CO2 delivered
at approximately 3 psi. The CO2 was delivered from a small tank
strapped to a pack frame for easy portability. A regulator maintained
constant flow and pressure from the tank through the rubber tube
to one of the glass tubes in a two hole rubber stopper inserted in
the vial. The other glass tube in the stopper served as an exhaust
to avoid excessive pressure in the vial ( Figure 3 ) • The anaesthetized
butterfly was removed from the vial with flat bladed insect forceps and
classified according to sex, forewing spot pattern and the area and
date of previous capture if it had been a recaptured specimen. The
specimen was then marked to indicate the date and area of capture.
This was done by putting a dot of "Pactra” lacquer on the wing
underside. The critical factor of this operation was maintaining a
proper paint consistency. This was done by trial and error, using
acetone as a diluent with a blunted dissecting needle. Six different
colors were used to denote the six stations. These were applied to
one of ten distinct underwing areas to denote the date of capture
(Figure 4). The butterfly was then carefully laid on the bottom of
a one gallon ice cream carton and allowed to recover. Recovery time
varied, but seldom exceeded two minutes. The process was then
repeated. After handling the last specimen, all the gear was assembled.
Just prior to moving to the next station the carton was held upright,
facing the sun, and was gently tapped so that the remaining specimens
would fly off. If any specimen remained, the carton was inverted and
vigorously tapped. If a specimen was not able to fly "normally” a
distance of 10 feet, it was removed from the population and the
event recorded. After the first day’s collection, subsequent collecting
within about 10 feet of the center of the station was avoided as
a precaution again recapturing injured specimens, if any should
exist. In 1955, 21 individuals including 10 recaptures, and in 1956,
37 individuals including 8 recaptures, were removed from the popula-
tioa These figures are not wholly indicative of the effectiveness of
the technique since about half of these represented specimens sampled
for study.
(4) .-237-244. 19^3
MATTONI AND SEIGER
241
Fig. 4. Marking scheme on underwings to denote data of capture. The
circles indicate the positions of the lacquer marks on the wings which corres-
pond to the collecting day. (designated by Roman numerals).
Fig. 5. Recaptured specimens sampled on the last collecting day of 1955.
242
PHILOTES SONORENSIS
/. Res. Lepid,
The distinctive feature of our technique is the use of COq anaes-
thesis in the field. We feel that anaesthesis greatly decreases the
probability of damage due to handling, especially when examining
and marking such a small butterfly as P. sonorensis. The procedure
was adopted because of its general acceptability in the laboratory
for research in insect physiology and genetics. For example, Seiger
(1953) studied the effects of different anaesthetics on the dipteran,
Drosophila mulleri. He found no significant difference in fertility and
fecundity between flies which had been given no anaesthesis and
flies which had been anaesthetized with COq. The average quiescent
period after anaesthesis was one minute. This was sufficient time
to classify an individual for sex and wing pattern, determine if it had
been recaptured, mark the lower wings, and allow the lacquer to
dry. Following recovery, the individuals tended to remain quietly
in the container, thus further minimizing damage. The lacquer fastens
the scales to the wing membrane and dries so quickly that no more
than ten losses were suffered by insects sticking to the container or
themselves. In the laboratory, P. sonorensis could withstand COq
anaesthesis in excess of 10 minutes with no apparent ill effects, although
under longer periods of exposure, partial paralysis would occur and
eventually death ensue. There was no evidence for cumulative effect
of repeated exposures of 10-30 seconds duration to COq. The possibility
that abnormal behavior might result from anaesthesis has not been
fully explored. In the first flight after anaesthesia, there appears to be
a tendency for the butterflies to exhibit an escape behavior. After alight-
ing once the behavior is not apparent. There appears to be no difference
between the behavior of a butterfly in its first flight after anaesthesis
and the behavior of a butterfly in flight after being captured in a
net and released without anaesthesis. Although we feel that the advan-
tages of anaesthesia far outweigh any possible disadvantages, we plan
to determine whether there are any real effects of COq on the behavior
of P. sonorensis in future experiments.
We believe that the most important evidence of the negligible effect
of our overall technique on behavior lies in the consistency of our
data for two years with respect to the highly non-random pattern of
movement. Another evidence was the remarkable behavior of 5
individuals in 1956. These moved away from their area of capture
and subsequently returned. There are several reasons which indicate
that viability effects are also negligible. Figure 5 shows four marked
specimens sampled on the last day of 1955. These appear to be quite
undamaged, that at the upper right having been followed for eleven
days and caught four times. In the course of our studies, several
insects flew the distance between the two farthest stations (433
meters), one flying a minimum of 819 meters. Lastly, because many
specimens were captured more than once, it is possible to infer
whether multiple handling had an additive effect on viability by com-
i(4)is37”244. 19^3
MATTONI AND SEIGER
243
paring observed with expected values of multiple recaptures 0, 1, 2, 3
and 4 times.
Using a poisson distribution, the data for 1955 gives = 23.7
and P = < .0001 for 3 df; and for 1956, = 5.4 and P = .12
for 3 df. The highly significant departure from expected in 1955
is due entirely to an excess of specimens recaptured 3 and 4 times. The
meaning of this departure is obscure, although it does not contradict
the hypothesis of additive deleterious handling effects.
We found that the marking technique enabled us to distinguish
individuals without confusion. The method can be compared to a
punch card system in which each butterfly carries a recorded code on
its wings, the color of the spot indicating the station where each
capture took place and the position of the spot on he wing indicating
the date of each- capture. Thus the population size for each station can
be determined for any given collecting day (Dowdeswell, Fisher and
Ford 1949, Ford 1951), patterns of movement can be discerned and
the life span of individuals can be calculated. These characteristics can
be further quantified by correlation with maculation type and sex of
the individuals. These data and conclusions are being prepared for
publication in this journal
For comparative information on field techniques in population
study with Lepidoptera, the reader is referred to Abtot ( 1959), Ehrlich
and Davidson (I960), Evans (1955), and Fales (1959); as well as
those of Ford and his associates (op. dt.) We feel our technique
has the least effect on viability and behavior differentials, particularly
with reference to the use of anaesthesis; minimizing number of
identifying marks on each individual; limiting individual contact to
net, vial and forceps; and no more than 30 minute retention in the
field. The last item, of course, is part of our experimental design. We
feel that all such studies should take this factor into account, in spite
of statistical difficulties in treating same day recaptures.
We gratefully acknowledge the contributions of Mr. Jack Roper
of North American Aviation in providing the photographs for Figures
1, and 3, Dr. David Goodchild, CS & IRO for Figure 2, and Mr.
Roy Pence, UCLA, for Figure 5.
SUMMARY
1. A method for anaesthetizing and marking individuals in order to
determine population struaure is described.
2. The benefits and possible disadvantages of the method are ex-
amined and some applications of the technique are mentioned.
244
PHILOTES SONORmSIS
J. Re$. Lepid.
REFERENCES
ABBOTT, W. 1959. Local Autecology and Behavior in Pieris protodice. Bois-
duval and LeConte with some comparisons to Colias eurytheme Bois-
duval, Wasmann ] Biol. 17: 279-298.
DOWDESWELL, W. H., R. A. FISHER and E. B. FORD 1949. The
Quantitative Study of Populations in the Lepidoptera. Heredity. 3: 67-84,
ERHLICH, P. R. & S. E. DAVIDSON I960. Techniques for Capture —
Recapture Studies of Lepidoptera populations. Lep. News. 14: 227-229.
EVANS, W. H. 955. Retrieving marked Anthocharis reakertii. Lep. News.
9: 118.
FORD, E. B. 1951. The Experimental Study of Evolution. Australian and
New Zealand Association for the Advancement of Science. 28; 143-154.
FALES, J. H. 1959. A Field Study of the Flight Behavior of the Tiger
Swallowtail Butterfly. Ann. Ent. Soc. Amer. 52: 486-487.
SEIGER, M. B. 1953. The Effect of X-Ray Dosage on Fertility, Fecundity
and Incidence of Chromosomal Aberrations in Drosophila mulleri. Master’s
Thesis, University of Texas.
Journal of Research on the Lepidoptera 1 (4) ;245-248, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 196}
EARLY STAGES OF A SOUTHERN CALIFORNIA
GEOMETRID MOTH, DREPANULATRIX
HULSTI HULSTI (DYAR)
JOHN ADAMS COMSTOCK
Del Mar, California
Dr. Dyar, in 1904, named "Catopyrrha hulsti” from a specimen
contained in the Hulst collection under another name, without desig-
nating the locality of its origin. Dr. Rindge (1949) gave valuable
data on the species, and its two subspecies, D. carneolata B. & McD.
(1917) and D. verdiaria Grossbeck (1912) in his Revision of the
Genus. He lists D. carneolata as an Arizona form of Drepanulatrix
hulsti, and mentions California examples that seem indistinguishable
from Arizona specimens.
He places Drepanulatrix rindgearia Sperry (1948) as a synonym of
D. hulsti. He also notes that the early stages are unknown.
D. hulsti hulsti ranges throughout the greater portion of coastal
California, wherever its food plant, Ceanothus, occurs.
It flies most abundantly from May to August, but has been reported
as early as January, and late into September.
Eggs of this species were obtained this year (1962) in Del Mar,
on several occasions, from May 25 to late August. For the most part
these were laid singly on their sides. When first laid they were light
green, with a pearly luster. Shortly they changed to cream or light tan,
with contrasting dots and irregular tortuous bands of dark pink to
salmon-red.
The characteristic from is ovoid, with a flattened base and rounded
tip, but some approach a barrel-shaped form with flattened base. Our
illustration (figure 1 A to C) shows the extremes of this variation,
and also the early and late changes in markings.
Sculpture of the surface consists of longitudinal ridges, from 28 to
36 in number, running from the edge of the flattened base to the top.
These ridges have pearled’ edges, the pearls or raised points correspond-
ing to the numerous horizontal lines crossing between the ridges. The
raised pearls are small and relatively inconspicuous.
On the flattened base the ridges end abruptly, and within the circle
formed by their terminations the surface is covered by numerous
irregular hexagonal pits with raised walls, as shown in figure IB.
There is considerable variation in the size of eggs, the average being
1 mm. tall by 0.6 mm. wide. Eggs laid May 25 hatched June 3. The
larval egress was through one end, the remainder of the shell being
left intact.
245
246
JOHN ADAMS COMSTOCK
/. Res. tepid.
By comparison with the egg of Drepanulatrix monicaria it is larger,
the number of ridges greater, and the salmon-red markings heavier.
FIRST INSTAR LARVA: Length, 375 mm. Head width, 0.4 mm.
The head is wider than the first segment. The ground color is white
with a heavy spotting of black dots on a brown base each side of the
epicranial suture and the outer and upper two-thirds of each cheek.
There are two raised papillae on the front. The mouth parts are brown,
and the ocelli black on a white ground.
The first thoracic segment is narrower than the head, but wider
than the remaining segments. The ground color of all body segments
is white. There is a longitudinal middorsal irregular band of olive which
is somewhat restricted at the segmental junctures. A narrow irregular
dorso-lateral band parallels it. There are several rows of black papillae
running generally in a longitudinal direction. The placement of these
on the dorsal surface is shown in the illustration, (figure ID). Each
papillus is surrounded at its base by a white circlet, and mounts a
black seta at its tip.
The legs are hyaline, with a tinge of light yellow. The single pair
of prolegs and the anal prolegs are concolorous with the body.
LARVA OF 15 MM. LENGTH: Head width, 1.5 mm. Head, gray,
spotted with black, the spotting heaviest on the margin of the crown
and cheeks. The front is slightly less spotted. The mandibles and ocelli
are black.
The body ground color is gray. Most of the segments are crossed
transversely by five or six folds, on which are placed prominent black
papillae bearing black setae. The entire surface of the body is heavily
sprinkled with minute black and brown dots. These have a slight ten-
dency to form longitudinal lines. The spots on the ventral surface are
predominantly brown and are more definitely arranged in longitudinal
lines.
The prolegs are concolorous with the body, and the true legs are
less spotted and more hyaline.
MATURE LARVA: Length, 26 mm. Head width, 2.2 mm. Head;
color, gray-brown, heavily spotted with minute brown dots. The front
is finely ridged horizontally, with a line of six minute black dots running
transversely near the lower edge. The labrum is darker, and is ridged
longitudinally. The maxillae are dark brown, the ocelli black, and the
antennae translucent.
Across the center of each cheek is a lunate black band, beginning
near the epicranial suture, extending laterally, then arching inferiorly
to end near the ocelli. The setae are brown, and arise from black papillae.
The head is shown in front view on figure IE.
BODY: Ground color, whitish gray, nearly obscured by numerous
small brown and black dots. There is a suggestion of a double middorsal
longitudinal brown stripe in the thoracic area. A poorly defined dark
spiracular band, made up of dots, is present on some specimens. The
spiracles have narrow black rims and cream colored centers. The legs
(4) :245-248
JOHN ADAMS COMSTOCK
247
Figure 1
Early stages of Drepanulatrix hulsti hulsti. A. Egg, lateral aspect final
color phase, enlarged X 40. B. Base of egg, enlarged X 40. C. Egg. elongate
form, and early light phase, enlarged X 40. D. First instar larva, dorsal aspect,
enlarged X 21. E. Head of mature larva, front view, enlarged X 10. F. and
G. Mature larva, dorsal aspect, enlarged X 5.5. H. Pupa, ventral aspect, en-
larged X 6. I and J. Cremaster, ventral and lateral aspects, enlarged X 14.
Reproduced from water color drawing by the author.
248
GEOMETRID M.OTH
/. Res. Lcpid,
and prolegs are translucent, the latter being spotted with brown. The
crochets are red-brown, apparently biordinal in alignment, and developed
only along the lateral edge of the oval foot-pad, with short stubs only
on the medial edge. The setae are light brown, relatively short, and arise
from minute black papillae. ( figure 1 F and 1 G ) .
Pupation occurred on the floor of the rearing jar, among leaves and
debris, mixed with a few strands of silk.
PUPA: Length, 14 mm. Greatest width, through middle of wing
cases, 4.2 mm.
The anterior two-thirds of the pupa is relatively wide and plump.
The six terminal segments taper sharply to a distinctive cremaster, as
will be noted on figure 1 H. An enlargement of the latter is pictured on
figure 1 I.
The eyes are prominent, dark, and protruding. The cephalic end
is evenly rounded. The antennae and maxillae extend to the margins of
the wing cases. The spiracles are relatively small and nearly indisting-
uishable without a lens. There are no setae.
The color of the pupa is predominantly reddish brown, but the eyes
and cremaster are black.
The cremaster is pear-shaped, with the stem end narrowed and
elongated. Laterally it bears two globular bodies, one on each side, when
viewed in ventral aspect. Portions of the surface are nodular. The tip
bears several short spicules which arch dorsally. Two of these are
longer, and recurve ventrally. These details can best be grasped by
referring to the illustration on figure IH.
The texture of the body surface of the pupa is predominantly smooth
and glistening.
BIBLIOGRAPHY
BARNES, WILLIAM and J. McDUNNOUGH.,1917. Contributions to the
Lepidoptera. 3 (4) : PI. 28, Figs. 5-6.
DYAR, HARRISON G. 1904. A few notes on the Hulst collection. ?roc.
Ent. Soc. Wash. 6: 226.
GROSSBECK, J. A. 1912. Jour. N. Y. Ent. Soc. 20: 285.
RINDGE, FREDERICK H. 1949. A revision of the Geometrid moths
formerly assigned to Drepanulatrix. Bull. Am. Mus. Nat. Hist. 94 (5) :
233-298.
SPERRY, JOHN L. 1948. Three apparently undescribed Geometrid moths
from the southwest. Bull. So. Calif. Acad. Sci. Al (1): 6-7.
Journal of Research on the Lepidoptera 1 (4) :249-259, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A,
© Copyright 196}
THE EFFECTIVENESS OF DIFFERENT ISOTHIOCYAN-
ATES ON ATTRACTING LARVAE OF PIERIS RAPAE
WILLIAM HOVANITZ; VINCENT C. S. CHANG AND
GERALD HONCH
The California Arhoretum Foundation and the Lepidoptera Foundation,
Arcadia, California; The Los Angeles State College; and
The University of California, Riverside, California
The effectiveness of black mustard oil (allyl-isothiocyanate)
on attracting larvae of Pieris rapae at various concentrations has already
been described in this JOURNAL (Hovanitz and Chang, 1963).
This mustard oil is the commonly used oil for condiments and is pre-
pared by pressing from the seeds of black mustard {Bras ska nigra).
This oil is known also from various species of Cruciferous plants in
addition to black mustard. Other mustard oils are known from a wide
variety of plants, not all of them members of the family Cruciferae.
This paper describes some experiments on the selection by Pieris rapae
larvae on allyl-isothiocyanate produced by two different methods, and
on three other isothiocyanates.
NATURALLY OCCURRING ISOTHIOCYANATES
Isothiocyanates (mustard oils) are naturally derived from gluco-
sides which occur in a wide variety of the higher plants, belonging to
a relatively small number of plant families (Kjaer, I960). These
glucosides are characterized by the ability to undergo enzymatic hydro-
lysis to isothiocyanates, hyrogen sulfate and D-glucose. The latter
has invariably been encountered as the sugar moiety of the more than
thirty individual compounds recorded thus far (Kjaer, I960). Accord-
ing to this author, only nine glucosides of these thirty isothiocyanates
have as yet been crystallized. The occurrence of more than one gluco-
side in a given plant species is most common; as many as eight in-
dividual glucosides have been distinguished in a single seed specimen.
The compounds appear to be distributed over the entire plant; the
glucosides are diffusely present in parenchymal tissues, especially in
^ Aided by a grant from the National Science Foundation, Washington, D. C.
249
250
HOVANITZ, CHANG AND HONCH
/. Res. Lepid.
the bark (Guignard, 1890, 1893). The embryo constitutes the site of
accumulation in seeds. Little has been done to relate the variation in
glucoside content as a function of the stage of growth, or of environ-
mental factors such as climate, soil, etc, Stahmann, et al (1943) found
that 2-phehylethylisothiocyanate, the aglucone of gluconosturtiin, to be
the predominant mustard oil enzymatically liberated from the roots of
Brassica nigra, whereas the seeds of this plant represent the classical
source of the glucoside sinigrin which yields allyl isothiocyanate upon
enzymatic fission. Delaveau (1958) has noticed considerable varia-
tion in the total and relative amounts of the individual glucosides in
the plant Alliaria officinalis during its growth cycle. Considerable vari-
ation has been detected in quantitative and qualitative differences in
glucoside content of various parts of a plant as well as in different
lots of the same plant tissue.
A list of the known natural isothiocyanates and their parent gluco-
sides in various plant genera and species which are known to be hosts
of species of Pieris is given in table 1. In this table the plants are listed
by families and genera. After each genus or species of plants is listed
( 1 ) the Pieris known to feed on it, ( 2 ) the mustard oil glucoside
which has been isolated, or established as having been found, in that
plant, (3) the aglycones (the mustard oils corresponding to the gluco-
side) and (4) the group of compounds to which it belongs.
Observation of this list does not seem to indicate any common
denominator between the mustard oil, or glucoside, and the species of
Pieris which is attracted to it, with a single exception. That is, sinigrin
is present in most plants attracted by one or more species of Pieris.
This, in itself, may mean little for sinigrin appears to be the most com-
mon mustard oil glucoside, and the earliest known. There are Pieris
attracted by plants in which sinigrin is not known, for example,
( 1 ) Lepidium, which is a common food plant of Pieris protodice and
P. occidentalis, (2) Cleome (and probably Isomeris), which is a com-
mon food plant of P. protodice, and P, beckeri, ( 3 ) Propaeolum, a food
plant of Pieris rapae, and (4) various Resedaceae, food plants of Pieris
daplidice. In each of these plants indicated, there are some other mus-
tard oil glucosides present, namely, glucotropaeolin and glucoraphanin
for Lepidium, glucotropaeolin for Tropaeolum, gluconasturiin, gluco-
barbarian, and glucotropaeolin for the Resedaceae, and glucocapparin
for Cleome (or Isomeris, Capparidaceae) .
It seems likely that more than one mustard oil attracts Pieris,
even of the same species. The possibility exists that there may be little
specificity for the kind of mustard oil, so long as some one isothio-
cyanate is present in the proper concentration. Except for Tropaeolum,
all listed food plants of Pieris rapae contain sinigrin in some part of
the plant. Only glucotropaeolin is known from Tropaeolum itself, yet
Pieris rapae survives on that plant after only a few generations of adap-
tation.
i(4):249-259, 1963
ISOTHIOCYANATES
251
TESTS OF LARVAE OF PIERIS RAPAE TOWARD
VARIOUS ISOTHIOCYANATES
The tests to be described here were made by subjecting larvae of
Fieris rapae from our laboratory strain, grown for several generations
on black mustard {Brassica nigra), to our testing device. This device
has been described previously (Hovanitz and Chang, 1963). It con-
sists essentially of a greenhouse flat, with filter paper moistened with
dilutions of mustard oils, and spaced equally around the periphery of
the flat. The larvae are set in the center and make their way to the
side of the flat, being attracted to one or another of the mustard oils,
at various concentrations.
For this experiment, tests were made of five different mustard oils
in concentrations from 10~^ to 10~'^. These five were:
( 1 ) benzyl-isothiocyanate : this is the aglycone of glucotropaeolin,
found in Cruciferae: Lepidium sp., Resedaceae, and Tropaeo-
laceae; Tropaeolum sp. Synthetic origin.
(2) phenyl-isothiocyanate: there is no natural aglvcone of this
mustard oil listed by Kjaer. Synthetic origin.
(3) phenethyl-isothiocyanate: this is the aglycone of gluconasturti-
in, found in Barharea, Brassica nigra. Brassica oleracea and
the Resedaceae. Synthetic origin.
(4) allyl-isothiocyanate: this is the algycone of sinigrin found in
many plants. Commercial synthetic.
(5) same as above, only commercial natural product from Bras-
sica nigra seeds.
Sources of the above mentioned mustard oils are described below.
General Procedure for the Preparation of Synthetic Mustard Oils
The following general procedure for the preparation of mustard oils from
various amines was provided by Professor Henry Klostergaard of the San
Fernando Valley State College. This method was used for the preparation of
the phenethyl-, benzyl-, and methyl-isothiocyanates used in this paner. All
procedures to be carried out at 0°C.
In a 2000 ml. Erlynmeyer flask, mix 0.07878 moles of the amine and
enough absolute ethyl alcohol to make a 30% solution ( by volume). To
the above solution add 4.8 ml. CS^ solution (0.03939 moles in a 1:1 solution
with absolute alcohol). Wait 10 to 15 minutes, stirring often as there will
be some heat evolved. Now add 100 ml. of the iodine solution (0.03939
moles or 10 gms. of iodine dissolved in 100 ml. absolute alcohol). The
solution will start to clear up. Disregard any crystalliztion at this point.
Shake until the brown color is gone and some yellow color appears. This is
not stable so immediately go on to the next step. At this point add 25 ml.
of the sodium solution (0.03939 moles or 0.9 gms. of sodium in 25 ml.
absolute alcohol). The mixture will split and become homogeneous.
Now add another 100 ml. of the iodine solution as above. Some ele-
mentary sulfur will settle out (this is not soluble in ether). Now add 250 ml.
acidifed water (1% HCL).
Next pour the solution into a separatory funnel and extract the mustard
oil with ether. The mustard oil will appear in the ether phase or the top
TABLE 1. The occurrence of natural isothiocyanates and their parent glucosides in plant
and species known to be hosts of species of Pieris (chemical data from Kjaer) .
252
HOVANITZ, CHANG AND HONCH
J. Res. Lepid.
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ISOTHIOCYANATES
253
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254
HOVANITZ, CHANG AND HONCH
/. Res. Lepid.
layer. Discard the lower layer. Add some 10% NaOH to the mustard oil
and ether solution to take off the iodine and then discard the lower layer.
Next wash the solution with an equal volume of distilled water. Discard
the water layer. Now there is present an ether solution of the mustard oil,
(Ice this solution immedately.) Vacuum distil this solution in a flash evap-
orator with the solution in the flask cooled by an ice-salt solution and suc-
cessive flasks cooled by ether and dry ice. Keep the mustard oil and ether
solution under vacuum for at least IVz hours. The mustard oil will be left
in the original flask. Remove it immediately and store at 0°C.
Caution
All the solutions involved in this preparation should be kept refrigerated
at 0°C. until used. All work should be done under an adequate hood since
the chemicals involved are highly toxic and odoriferous.
Tests of the larvae of Pieris rapae as indicated were made separately,
utilizing the five dilutions of the mustard oils indicated. The actual
dilutions may actually have been less than indicated because it appears
that at the temperature used (about 20 °C.), not all the mustard oil
may have gone into solution. Each larva was tested twenty times, giving
total test times ranging from 180 times for benzl-isothiocyanate to
680 times for phenyl-isothiocyanate.
The allyl-isothiocyanates differed in their response, according to
their origin. One was derived from black mustard seeds by standard
commercial techniques of compression, and fermentation. The other
was synthetically prepared. In each case, the larvae selected the 10“*^
dilution of mustard oil (22.1%, for synthetic to 21.2% for natural)
but the dispersal of the selections was different. For example, the
dilution of 10~^ was selected by the larvae for the synthetic mustard
oil 20.7% of the time, as compared with only 11.6% for the natural
product. Dilutions of less than 10~^ are also different. For example,
while the synthetic gave selections of 193% at 10~®, the natural
product gave 14%. At 10~'^, these were 15.2% for the synthetic as
compared with 11.6% for the natural. An apparently significantly
greater number of larvae left the flat without any selection at all with
the natural (24.2%) as compared with the synthetic product (2.5%).
This discrepancy was probably due to the human variation in carrying
out the tests. The percentage is greater when the larvae are not allowed
as long a selection period. This discrepancy would not affect the actual
selections themselves.
Another effect is quite noticeable with these data, as well as with
the data on the other mustard oils, namely, that the selections give a
bimodal curve (see fig. 1). Such a bimodal curve was not clear with
our previous testing though a trace of it might be detected (fig. 3,
Hovanitz and Chang, 1963). The previous testing was made with
the natural product which also shows a very poor bimodal curve ( fig. 1 ) .
The synthetic product shows a more strongly indicated bimodality at
10“'^ and at 10~® (fig. 1). The reasons for the bimodal curve which
is especially apparent for the synthetic allyl mustard oil are unknown.
Selections for benzyl-isothicoyanate are almost certainly negative.
TABLE 2. The selection by larvae of Pieris rapae for various concentrations
of mustard oils.
(4):249-259, 1963
ISOTHIOCYANATES
255
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256
HOVANITZ, CHANG AND HONCH
/. Res. Lepid.
25
Fig. 1. Histograms illustrating the fomparison between natural and snythetic mustard
oils respecting their selection by larvae of Pieris rapae. Note the strong bimodality of the his-
togram showing the synthetic product as compared with the natural
PERCEN T PERCEN T PERCEN T
(4);249-259, 1963
ISOTHIOCYANATES
257
-
—
BENZ)
YL
10- ®
10^
10'®
10-3
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NONE
f
Fig. 2. Histograms illustrating the selections made by larvae of Picris rapae for various
dilutions of the mustard oils benzyl-isothiocyanate, phenyl-isothiocyanate and phenethyl-ioso-
thiocyanate. Note the bimodality of two of the histograms and the nearly complete lack of
selection indicated by the other.
258
HOVANITZ, CHANG AND HONCH
/. Res. Lepid,
not varying significantly from water alone (table 2 and fig. 2). This
mustard oil also was the one in which dissolving in water appeared
almost impossible and in which case the tests might very well have
been conducted on water alone. Perhaps at a higher temperature, dis-
solving the mustard oil might have been accomplished but under the
circumstances it does not seem as if this mustard oil could be of any
influence on attracting Pieris rapae larvae to Tropaeolum species where
it is found naturally.
Phenyl-isothiocyanate and phenethyl-isothiocyanate were about
equally effective in attracting the larvae though there is a possibility
that the phenyl mustard oil may be slightly less attractive than the
phenethyl, since at each concentration the percent attraction was
slightly less. However, the decrease possibly is not significant. The
attraction of both of these mustard oils was not much different from
that of allyl isothiocyanate (table 2 and fig, 2). This leads to the
interesting possibility of the specificity between isothiocyanates in the
generic sense and the Pieridae that are attracted to plants that contain
the compounds. It is possible that the attraction is to the isothio-
cyanate part of the molecule and not to side chains. "When applied
to the tongue, all mustard oils cause a sharp and burning sensation.
Their odors, though mostly pungent, display characteristic individual
differences which often are helpful in the detection and classification
of mustard oils. Certain isothiocyanates, that undergo rapid intramole-
cular cyclization, give rise to a transient biting taste, followed by a
sensation of bitterness. Like most synthetic mustard oils, those of
natural origin show vesicant and frequently also lachrymatory proper-
ties,” from Kjaer (I960). These same substances, however, when
present in small concentration appear to give rise to the desirable con-
diment properties of the mustard oils and appear in turn account for
the specificity of these oils to the larvae and adults of Pieris.
CONCLUSIONS AND SUMMARY
1. A list is given of the mustard oils which are known to occur in
food plants of the Pieridae. These are derived from naturally-occurring
glucosides which are also listed.
2. It seems evident from this list that more than one mustard oil
attracts Pieris, since the evidence does not indicate that only one could
be responsible.
3. Tests were made on the selection of larvae of Pieris rapae
toward several mustard oils at various concentrations. These oils were
either commercially obtained, or were synthetically compounded by us.
The procedure used for synthetic compounding is described.
4. Synthetic and natural allyl mustard oils obtained commer-
cially gave somewhat divergent testing results. The synthetic gave a
bimodal curve with modes at 10”^ and 10”® which were only slightly
(4)5249-259, 19^3
ISOTHIOCYANATES
259
or flot at ail apparent with the natural.
5. Other mustard oils tested were benzyl-isothiocyaoate, phenyl-
isothiocyanate and phenethyl-isothiocyanate.
6. Benzyl-isothicoyanate is the aglycone of glucotropaeolin, the
glucoside of Tropaeolum sp. and some other plants. Its attraction power
at the concentrations used is not different from that of distilled water.
In view of the difficulties experienced in dissolving this substance^ it
is possible that none actually dissolved and that, therefore, the tests were
actually on water alone.
7. Phenyl-isothiocyanate is not known to be natural occurring.
Nevertheless, it had a selective effect on the larvae only slightly less
effective than allyl- and phenethyl-isothiocyanates.
8. Phenethyl-isothiocyanate had an attractive power almost equiv-
alent to that of the synthetic allyl-isothiocyanate. It is a naturally-
occurring mustard oil, being the aglycone of gluconasturtiin which is
found in Nasturtium and Barbarea.
9. It appears that more mustard oils than just allyl-isothiocyanate
are attractive to Pieris. The experience with benzyl-isothiocyanate
however indicates that not all are effective, at least not under the ex-
perimental conditions utilized.
LITERATURE CITED
DELAVEAU, P. 1958. Variations de la teneur en heterosides a senevol de
VAUiaria officinalis L. an corns de la vegetation. C. R. hebd. Seances
Acad. ScL 246: 1903.
GUIGNARD, L. 1890. Recherches sur la localisation des principes aaifs des
Cruciferes. /. Botanique 4: 385.
1893. Recherches sur la localisation des principes actifs chez les
Capparidees, Tropeolees, Limnanthees, Resedacees. /. Botanique 7: 345.
HOVANITZ, W. and V. C. S. CHANG. 1963. Selection of allyl-isothiocyanate
by larvae of Pieris rapae and the inheritance of this trait. /. Res. Lepid.
1(3) : 169-182.
KJAER, ANDERS. I960. Naturally derived isothiocyanates (mustard oils)
and their parent glucosides. Fortschritte d. Chem. org. Naturst. 18:
122-176. Springer, Wien.
STAHMANN, M. A., K, P. LINK and P. C. WALKER, 1943. Mustard oils
in crucifers and their relation to resistance to clubroot. /. Agric. Res.
67: 49.
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Journal of Research on the Lepidoptera
1(4) :261-274, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 1^6}
THE ORIGIN OF A SYMPATRIC SPECIES IN COLIAS
THROUGH THE AID OF NATURAL HYBRIDIZATION
WILLIAM HOVANITZ
The Lepidoptera Foundation and the California Arboretum
Foundation, Inc., Arcadia, California and the Los Angeles
State College, Los Angeles, California
1. INTRODUCTION
The theoretical set up for the origin of a new species in any
biological group is the existence of two or more species in the same
geographical area, which mix and reassort their combined gene com-
binations by the process of hybridization, Fi intercrossing and back-
crossing. There have been examples described in recent years in which
crossing of this sort has served to blend together two existing species
in a given area in such a way that both parental species cease existence
as independent entities; this situation has been described generally as
"subspecific intergradation” because of necessity the blending must be
done on a geographical, or spatial, scale. Recent examples of this have
been described by Sette (1962) involving the subspecies macaria and
laufina of Argynnis callippe, and others by Hovanitz (1949).
In those cases in which natural hybridization of the two species has
served only to permit gene (and character) interchange between the
two original species, both of which maintain themselves as independent
entities, but in which no new species is formed, the situation is described
as one of introgressive hybridization. One of the most extensive cases
of introgressive hybridization known is that existing between the
species Colias emytheme and Colias philodice (Hovanitz, 1943, 1944a,
1944b and 1948). Other cases which are less extensive are known in
many other species of Colias.
Much more rare, possibly because the occurrence may be transient,
is the situation in which two hybridizing species retain their separate
existence, but out of the assorted gene pool arising by hybridization,
there come a third species which maintains itself as a separate entity
without the further necessity of hybridization. This is the situation to
be analyzed in this paper.
It is almost axiomatic that for closely related and interfertile species
to exist in the same general area, and yet to maintain their separate
identities, those two or more species must be separated in some way
261
262
HOVANITZ
/. Res. Lepid.
Fig. 1. North polar projeaion map showing the world distribution of
Colias hecla.
hy physiological or ecological isolation barriers for if this were not so,
the genes of the two or more species would be blended together as in
one gigantic melting pot. The isolation barriers for Colias appear to be
such physiological and ecological ones rather than genetic ones since
most species appear to be interfertile. Tied with these barriers is the
selection for genes controlling adaptation for different habitats and dif-
ferent food plants of the larvae, which enable Colias to exist under a
greater variety of conditions than would be possible were they all
amalgamated into one species in a given geographical area.
Previous study has indicated that there is considerable natural
hybridization in the genus Colias (Hovanitz, 1949, 1956). The species
Colias philodice and Colias eurytheme have been shown to hybridize
rather freely throughout all parts of their ranges where they are in
i(4):i6i-274. 5963
SYMPATRIC SPECIES
263
\ ■ ■ —I-- , i ■■ ■ »■ ' ^ Colios nQSt0S l-AWBERT'S AZIMUTHAL EQUAL-AREA PROJECTION
Colias phicomone
Fig. 2. North polar projection map showing the world distribution of
Colias nastes.
contact, or where there is overlap of their habitats (Hovanitz, 1944a,
1944b). This is a classical example of intro gression, the reciprocal
transfer of genes between two or more species in natural contact, but
without the complete submergence of the identity of any. Similar
examples of natural hybridization have been described for the following
species combinations of Colias:
Colias interior - C. christina
Colias hyale - C. erate - C. croceus
Colias Christina - C. gigantea
Colias philodice = C. eurytheme
Colias hecla - C. nastes
Colias interior - C. philodice
Colias alexandra - C. philodice
Colias Christina - C. philodice
264
HOVANITZ
/. Res. Lepid.
Fig. 3. North polar projection map showing the world distribution of
Colias palaeno.
Without doubt a great number of additional combinations will be
found, wherever any two or more of the species of this genus occur
together. Species hybridization in the laboratory, or in the field, lead-
nig only to Fj generation individuals, with no history of fertile back-
crossing or of fertile Fo production, would have no effect on altering
the composition of natural populations. Such crosses therefore are
only indicative of physical ability to mate, of Fi to survive (though
not of Fi fertility), and to indicate something of the nature of genetic
dominance of the many characters involved in the cross; however, they
give no information at all on the evolutionary relationships between
the parental species. Further testing must be done for this purpose by
carrying the crosses through to the F2 generation and backcross gener-
ations, and by study of the genetics of the natural populations them-
selves.
(4):26i-174. 1963
SYMPATRIC SPECIES
265
Fig. 4. Distribution of Colias hecla in North America, together with
that of Colias meadi. The ranges of these two species do not overlap
but are separated at their closest point by an altitudinal difference of
4000 feet. Colias hecla is found in the mountains and plains of the
north and Colias m.eadi only in the mountains of the south.
266
HOVANITZ
/. Res. Lepid.
Pig. 5. Distribution of Colias nastes in North America. The species
is found only in areas north of the tree line throughout the eastern and
central parts of its range but occurs also far to the south in the western
region of the Rocky Mountains in the alpine tundra.
I (4) :i6i-274, 1963
SYMPATIUC SPECIES
267
It has been shown that Colias hecla and Colias nastes give all indica-
tions of hybridizing in the wild. These indications are as extensive as
those which are known for Colias eury theme and Colias philodice hy-
bridizations, with the exception of a lack of the actual breeding ex-
periments. In other words, the information is available by an analysis
of wild populations, and character intergradation within the popula-
tions. It has been conjectured, in other publications, that these two
species have hybridized in a certain area, namely the Canadian eastern
arctic, in such a way and to such an extent, that a third species, Colias
boothi, has originated as a result of such hybridization (Hovanitz,
1949). In the areas concerned, both parental species and the "inter-
mediate” newly originated species all survive without the complete
submergence of either parental species, or of the new species. The
evidence indicates that on the fringes of the distributional area occu-
pied by the "new species,” intermediates are present in a way suggest-
ing introgression only, rather than the independent existence of the
new species. This "origin of a new species by hybridization” has been
of theoretical significance in problems of evolution for many years;
the present example is the first clear authenticated case of this type
of origin which can be observed in action.
II. GEOGRAPHICAL RELATIONSHIPS BETWEEN COLIAS
HECLA, COLIAS NASTES AND COLIAS PALAENO
The three most "northern” Colias in the world are Colias hecla,
Colias nastes and Colias palaeno, the most northern of these three
being first and palaeno being least northern. Although only the first
two appear to be involved in the natural hybridization relationship, the
third is considered here because of its frequent sympatric geographical
distribution.
The world distributional ranges of these three species are shown in
figures 1, 2, and 3. The North American ranges are shown in more
detail in figures 4, 5 and 6. The latter three maps have been pub-
lished previously (Hovanitz, 1951).
Colias hecla is distributed from northern Norway and Sweden
across the arctic shores of Asia ( fig. 1 ) , across the Bering straits, across
Alaska and northern Canada, throughout the Canadian arctic islands
and around all coasts of Greenland. Southern extensions of range
occur in several places around this circumpolar range. In Scandinavia,
the species extends part way down the Sandinavian peninsula, but it
does not appear in more southern European locations. In Asia, besides
the coastal arctic locations, the species exists inland in the Verkhoyansk
area and along the Lena river valley. Farther southwards, there is much
material indicating the possibility of its existence in almost all ranges
of central Asia including all the area from Hindukusch, Pamir and
Karakorum across the Himalaya to Ladakh and mountains east and
268
HOVANITZ
/. Res. Lepid.
north. The ranges extending north and eastward from the Pamir to the
Baikal and Transbaikal completes a circle of mountains around the
Gobi desert where some form of Colias hecla appears to live. Dots
marking the exact locations for this area on the map were omitted
because of the difficulty in every case to be certain of the identifications
between the hecla group of forms and a group related to Colias meadi
of the mountains of North America. Both of these groups appear to
be in the same areas of central Asia, though they are separated in North
America into contiguous regions.
In North America, Colias hecla is found in Alaska in all areas ex-
cept the parts southern and coastal of the main southern mountain
masses. This includes the mountains and valleys of the interior, and
the western and northern coastal plains. In the Rocky Mountains, the
species is found as far south as the present location of the Alaska
highway from Whitehorse to Ft. Nelson but there are no locations
known south of this point in the mountains area. The species then
extends southwards on the Rocky mountain piedmont and the valley
of the tributaries of the Mackenzie River as far south as Nordegg (just
west and south of Edmonton, Alberta). The range skirts the prairie
country of central Canada near Lake Athabaska, eastward to Fort
Churchill on the Hudson Bay. The species is found on the most north-
ern part of the Ungava peninsula and the upper tip of Labrador, but not
along the Labrador coast. The map (fig. 4) in addition to showing
the distributional range of C. hecla also shows the range of C. meadi,
the relationship of which to C. hecla is uncertain despite its close
phenotypic appearance.
Colias nastes has a distributional range only slightly more "southern”
than that of Colias hecla. Actually, the ecological requirements of C.
nastes, besides a temperature generally below 50 °F., are somewhat
different from that for C hecla as will be seen later. Unlike C. hecla,
Colias nastes does not exist on any part of Greenland or the most
northern of the Canadian arctic islands. Thus, on its circumpolar
range, there is a break of great proportions across the Atlantic which
does not appear on the map for C. hecla. Colias nastes is found in
Scandinava in the same general locations as Colias hecla, extending part
way down the mountain range. From there, it extends across the
European and the Asiatic coastal sections (presumably) across the
Bering straits to Alaska, along the north coast of Alaska and Canada
to Victoria Island and Baffin Island but not to the islands to the north.
In the islands north of Europe and Asia, C, nastes is known in Novaya
Zemlya and has been listed as having been taken in Spitzbergen. South-
wards, the species as such is not known in Europe, unless Colias phi-
comone of the Alps and the Pyrenees should be considered the same
species. There seems little reason for not so considering it, in view of
the fact that they are so similar in most respects. However, if they
were connected in their distributional range at some earlier time, as
(4):i6i-274. 1963
SYMPATRIC SPECIES
269
Fig, 6. Distribution of the Faccmiwm-feeding Colias of North Amer-
ica. These include Colias palaeno of the mountains and plains of the
northwest, Colias interior of the plains in the southern part of the
range as well as the Cascade and Appalachian Mountains, Colias pelidne
of the eastern arctic and the northern Rocky Mountains, and Colias
l)ehri of the alpine regions of the Sierra Nevada.
iisiHr . j ' \ f
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iamiiiMi»aiJiamaiBiaiatatnatnnBiBiaiaiaiataiBinaiaiaiaiBiaMaiaaai«i«i5iSi5M«S5i5
ia|a|ii|g||||||H|^ataiaiaiaiBiaiBiataiaiaiiiiii|I|||i|i^^aiaiaiaiaiaiaiaiaiaiStaia
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Fig. 8. Series of Coiias, mostly identifiable as Colias boothi, showing
by means of a graded series, nine steps in the border pattern transformation
from the male (or hecla) pattern at the top left to the female (or nastes)
pattern at the second to the bottom right. Grades are designated as 8 through
4 on the left column and 3 through 0 on the right column. The specimen on
the bottom right does not enter the series but is shown to illustrate a
weak pattern development on a grade 0 male. All specimens shown are
males and all are from Coppermine, Northwest Territories, Canada, July 12-
19, 1947, W. Hovanitz coll, except: (1) 2rd from top on right column:
Repulse Bay, Northwest Territories, Canada. July 21, 1950. P. F. Brugge-
mann. (2) 5th from top (last one) in right column. Repulse Bay, July 15,
1950. P. F. Bruggemann.
272
HOVANITZ
/. Res. Lepid.
during the Pleistocene ice age, many thousands of years of isolation
since that time has brought differences between them and earned a
species distinction. Southward in central Asia in the ring of mountains
surrounding the Gobi desert, including the Himalayas there is a string
of races all of which appear to be specifically related to C. nastes; the
main form here is known as C. cocandica. The map for C. nastes
therefore has quite a similarity to the map of C. hecla. It can be seen
however that in the area of Asia west of Sakhalien Island, C. nastes
occurs considerably south of the area of C. hecla.
In North America, Colias nastes occurs also farther south than C.
hecla. The species extends southwards in the Rocky mountains from
the mountains of Alaska and the Yukon Territory to the borders of
the United States in British Columbia and Alberta. In Alberta, C.
nastes occurs at an elevation of 7500 feet and up while C. hecla is
found at about 3-4000 feet in the piedmont. In these areas, C. nastes
is flying in late July and August while C. hecla at the lower elevations
is flying in May. C. nastes does not extend into the lowland valley areas
of the Mackenzie tributaries as does C. hecla but instead is found only
to the north of the tree line from the mouth of the Mackenzie to the
area of Ft. Churchill on the Hudson Bay. It is found on the Belcher
islands in Hudson Bay. It is found in the northern part of the Ungava
Peninsula and then southeastwards along the Labrador coast half way
down. The fost northerly known locality is on the northern coast of
Baffin island. These locations may be observed in relation to one
another on the map (fig. 5).
Though Colias palaeno is not involved in the specific study here
being analyzed, it was thought that it might be so involved and therefore,
its geographical distribution is here given together with the two pre-
ceding species. This species has a generally more southern distribu-
tional range than either Colias hecla or Colias nastes as can be seen by
study of the world map (fig. 3). The species is found in the Alps of
Europe, in a number of locations in the territory intervening between
the Alps and the Scandinavian mountains, and northwards and east-
wards. It does not exist far south of Lake Baikal in central Asia but
does extend southwards into Northern Korea, on the island of Hondo
in Japan, Sakhalien Island, and Kamtchatka Peninsula. The species is
not known on the islands north of Europe and Asia. In North Amer-
ica, the species is known in Alaska south of the Brooks range and north
of the main southern ranges which skirt the coast. It reaches the coast
of Northern America only at the mouth of the Mackenzie river but
extends southwards along the route of the Alaska highway to a point
just north of Ft. St. John and thence eastwards to the Riding Mountains
of Manitoba and northwards to the tip of the Ungava Peninsula. Its
range is limited southwards and eastwards by contact with Colias pelidne
(also called interior). For details on this contact zone, the map of
North America is more precise (fig. 6).
1963
SYMPATRIC SPECIES
273
IIL CHARACTERISTIC DIFFERENCES BETWEEN COLIAS
HECLA, COLIAS NASTES AND COLIAS PALAENO
Colias hecla may be distinguished readily from Colias nastes by
the fact that it is always orange in wing color (fig. 7), while Colias
nastes is pale yellow. Colias palaeno is more generally a brighter lemon
yellow (fig. 7).
A second character that separates hecla from nastes but not from
palaeno is a male-female dimorphism of the pattern. The male pattern
is a single solid band of black on the outer edges of the fore and hind
wings. This appears on the wings of the males of both hecla and
palaeno (fig. 7). The female pattern on the other hand dif feres from
this in a way that might be described as a series of dots in the border
band, or in what might be the band when it is all present. This is
typically shown for the females of hecla and nastes on figure 7. The
band of the female of Colias palaeno is so reduced that the dots show
only slightly in that species. The male of Colias nastes differs from
any other North American species of Colias in that it also shows a
series of dots in the border band, much like the female. In some
cases, it is very difficult to distinguish the males from the females by
general appearances.
Other differences in habit, larval food plant and ecological prefer-
ences will be discussed later in this series.
Intergradation between Colias nastes and Colias hecla as would be
caused by hybridization, the crossing of Fi to obtain the F2 segregation
and backcrossing should give a series of intermediate products with
all intergradations from the one parental type to the other. In addi-
tion there should be produced, if the genetic segregation truly assorts
itself with slight effects of genetic linkage, into truly divergent types
such as a fully orange male with the female border band, a type found
nowhere under natural conditions in the ordinary range of the species.
Another type of extreme divergence which would be found only under
such conditions of gene assortment would be a yellow form with the
typically male border pattern. All these types have been found in
areas of presumed hybridization of these two species, Colias hecla and
Colias nastes. A series of variations ranging from the male border band
to the female border band in a series of nine steps (grades) is shown
in figure 8. A similar graded series of nine steps for the orange pigment
on the upper side of the wings is shown in figure 9. These series are
to be used to illustrate the analysis of various populations of Colias
from diverse regions in the arctic.
274
HOVANITZ
/. Res. Lepid.
LITERATURE CITED (I, II, III)
HOVANITZ, WILLIAM. 1943. Hybridization and seasonal segregation in
two races of a butterfly occurring together in two localities. Biol. Bull.
85: 44-51.
— — — ■ 1944a. The ecological significance of the color phases of Colias
chrysotheme in North America. Ecology 25: 45-60.
— — 1944b. Genetic data on the two races of Colias chrysotheme in North
America and on a white form occurring in each. Genetics 29: 1-30.
— — -- 1948. Ecological segregation of interfertile species of Colias. Ecology
29: 461-469.
1949. Increased variability in populations following natural hybridiz-
ation. In ’'Genetics, Paleontology and Evolution,” Princeton University
Press, pp. 339-355.
— - — - 1951. The biology of Colias butterflies. I. The distribution of the
North American species. Wasmann J. Biol. 8: 49.75.
1956. Hybridization and species blending in the butterfly genus
Colias. Proc. XIV Int. Congr. Zool. Copenhagen, 1953, p. 140.
SETTE, O. E. 1962. Variation in the silvering of Argynnis {Speyeria) callippe
in the interior m*ountain area of southcentral California. /. Res. Lepid.
1(1): 3-20.
(to be continued)
Journal of Research on the Lepidoptera 1 (4) :275-279, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S,A.
© Copyright 196}
A METHOD FOR BREEDING PIERIS NAPI
AND PIERIS BRYONIAE
BJORN PETERSEN
Zoological Institute, University of Lund, Lund, Sweden
Since the year 1942 the author has bred Pieris napi L. and
Pieris hryoniae Ochs, over a number of years. This breeding was carried
out mainly in order to obtain information on the genetical, ecological,
and taxonomical relationships between the two forms (cf. Petersen
1963, in press, and literature cited there). This paper will deal with
the breeding methods used and some results obtained in this connection.
MATING
For copulation males and females were put together in cylindrical
cages made of cotton net with a bottom of treetex. The cages were
2-4 dm high and had a diameter of 2-3 dm. In such a cage 2-4 females
and a slightly larger number of males were placed. The cages were
hung up in a window facing towards the south. Quite soon it was
obvious that mating propensity was higher for napi in sunshine and
for hryoniae in cloudy weather ( cf . table 1 ) . Mating propensity means
in this case the frequency of maxing of one sex compared with the
same sex of the other species. The difference is statistically significant
(X^=l6.37 with Yates’ correction; P<0.001). Muller and Kautz
(1938 p. 15) state that sunshine is essential for the copulation of
hryoniae. In nature hryoniae, like napi, copulates in sunshine. The
experiments indicate only different ecological amplitudes for the copula-
tion of the two species.
Matings under laboratory conditions were fairly easily obtained, not
only by the author in Sweden, but also by Bowden and Easton in Eng-
land (Bowden 1953, 1956, 1957, Bowden and Easton 1955). Ento-
mologists in Central Europe, on the other hand, have had a number of
difficulties in obtaining such matings. Some few experiments on hybrid-
ization have been reported by Fischer (1924, 1925) and Kautz (Muller
and Kautz 1938 p. 159-161). The lack of success in many of the
Central European experiments may be due to too high temperature
during sunshine in Central Europe. The temperature in the windows
used by the author, when the experiments were made in April and
May, was usually about 24°-27°C. In the sunshine of Central European
laboratories the temperature probably is higher, especially as the ex-
periments usually are carried out during the summer months.
275
276
PETERSEN
/. Res. Lepid.
TABLE 1, Number of matings (f) of animals tested on
mating propensity (m) under different conditions. Sim-
plified from Petersen and Tenow, 1954, p, 182,
sunshine
cloudy
weather
f
m
f
m
napi
31
0,84
1
0,14
bryoniae
6
0,16
6
0,86
napi
22
0.76
2
0,29
bryoniae
7
0,24
5
0,71
TABLE 2, Survivals from growing P* bryoniae from
Abisko, northern Sweden,
temperature
number of
eggs
larvae
pupae
40
10
0
0
80
10
3
0
13°
10
9
6
16°
10
10
9
20°
20
20
11
28°
10
7
0
0
CM
00
22
14
0
0
=i-
CO
5
0
0
(4)s275-.i79. 1963
BREEDING PIERIS NAPI
277
EGG-LAYING
After copulation, the female was transferred into a cage with some
flowers to feed on and leaves of a species of the family Cruciferae for
egg-laying. P. bryoniae in nature lays the eggs almost exclusively on a
single plant species, Biscutella laevigata. In one cage experiment a
female preferred some other plants to Biscutella, about 80 eggs being
laid on them and none on Biscutella ( Petersen l.c. ) . Thus, every plant
appropriate for the egg-laying of napi seems to be appropriate for
bryoniae. However, most plants of families other than Cruciferae as
well as some few species of this family are not eaten at all by the larvae.
TABLE 3. Results from breeding napi - bryoniae from four
localities in Sweden, 1. = number of newly hatched larvae,
p, s number of pupae.
temp.
Scania
Uppsala
Murjek
Abisko
1,
P*
%
1,
P*
%
1,
P«
%
1,
P*
%
11®-12°
30
11
37
10
2
20
15
0
0
15®-16°
22
7
32
25
19
76
11
1
9
16^-17°
85
46
54
5
5
100
4
3
75
24
12
50
190-20°
153
119
78
45
26
58
17
8
47
33
19
58
220-23°
43
29
67
5
2
40
36
11
31
250-26°
92
47
51
5
4
80
STERILITY
In many broods of the species as well as of their hybrids, the eggs
do not develop at all or only partly. Sterility appears with almost the
same frequency in both types of crosses, 6 broods of 30 in the pure
species, 6 of 28 in the hybrid crosses. Sterility is slightly more frequent
among Bowden’s Fi hybrids and backcrosses, but the difference is not
significant. The F2 and F3 crosses, on the other hand, have a significantly
higher frequency of sterility, when compared with the Fi crosses and
back-crosses of Bowden (x^ = 5.94 with Yates’ correction, 0.02 >P
>0.01).
278
PETERSEN
/. Res. Lcpid.
TEMPERATURE
The influence of temperature on the breeding results was investi-
gated in two series of experiments. In the first series, material from
Abisko in northern Sweden was used. P. bryoniae flies here from 400 m
up to 800 m in the subalpine and the lower parts of the alpine regions.
The breeding results are shown in table 2. The optimal constant tem-
perature for eggs and larvae was found to lie between 13° and 20° C.
As the hibernating pupae require a rather low temperature the experi-
ments were stopped at pupation.
In the second series, material from four Swedish localities was used
( cf. table 3 ) . The Abisko material developed fairly successfully from
newly hatched larvae to pupation at 16° -23° C. The material from
Abisko was perhaps a little weaker than that in the experiment already
described, as the eggs had to be sent by mail. The material from south-
ern Sweden (Scania) developed at temperatures varying from 11° to
26° C. The adaptation to different climates is obviously not strong
enough to show up in experiments on such a small scale, especially near
the upper and lower limits of temperature.
DISEASES
A successful breeding is dependent, not only on appropriate food
and temperature, but also on the health of the animals. Virus diseases
often cause the blackening and death of the larvae. The cultures started
first usually develop well, only some of the latest-developing larvae
being affected. In later cultures the disease spreads like a pest, killing
the larvae in great numbers. The results of table 3 may therefore better
be expressed as different resistance to the disease at different tempera-
tures, or perhaps as varying virulence of the virus, than as a direct effect
of the temperature.
Most years, the larvae were kept in glass jars on cut leaves. As the
disease seemed to spread more rapidly on wet leaves, it was decided
to start breeding on living plants growing in flower pots. The results
were promising in that the frequency of virus disease became very low.
The fecundity of the animals, on the other hand, was not very high
during this last year of more extensive breeding. It has been stated
by several breeders of Pieris napi that it is impossible to breed many
generations without crossing with fresh material taken in nature. The
animals used belonged to the fourth generation in captivity and were
unfortunately the only ones available at the time.
The best way of breeding Pieris napi has still to be investigated. '
Breeding at 20° C on a living plant of the family Cruciferae, which
is easy to keep in flower pots, will probably give better results than
any so far known. Biscutella laevigata, which grows as a weed in the
gardens of Italy, will perhaps be the best food-plant for P. bryoniae.
(4):275-279. 19^3
BREEDING PIERIS NAPI
279
LITERATURE CITED
BOWDEN, S. R. 1953. Timing of Imaginal Development in Male and
Female Hybrid Pieridae. (Lep.) Entomologist, 86, No. 11: 255-264.
1956. Hybrids within the European Pieris napi L. Species
Group. (Lep., Pieridae). Proc. South London Ent. Nat. Hist. Soc.,
1954-55: 135-159.
— , 1957. Diapause in Female Hybrids: Pieris napi adalwinda
and Related Subspecies (Lep.) Entomologist, 90, No. 1133: 247-254,
273-282.
BOWDEN, S. R. and N. T. EASTON. 1955. Diapause and Death: Further
Observations on Imaginal Development in Pieris Hybrids (Lep.) Ento-
mologist, 88, No. 1108: 174-178, 204-210.
FISCHER, E. 1924. Ober die Zweibriitigkeit der P. bryoniae O. Mitt.
Munchen. Ent. Ges., 14:8.
— . 1925. Neue Zuchtergebnisse bei Pieriden. Mitt. Schweiz.
Ent. Ges., 13-
MULLER, L. and H. KAUTZ. 1938. Pieris bryoniae O. and Pieris napi
L. Wien. 1- 189.
PETERSEN, B. 1963. Breakdown of differentiation between Pieris napi L.
and Pieris bryoniae Ochs, and its causes. Zool. bidr. Uppsala, 35: in press.
PETERSEN, B. and O. TENOW. 1954. Studien am Rapsweissling und
Bergweissling (Pieris napi und Pieris bryoniae). Isolation und Paarungs-
biologie. Zool. bidr. Uppsala, 30: 169-198.
Journal of Research on the Lepidoptera
1(4) :281-300, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright
AN ANALYSIS OF THE NORTH AMERICAN SPECIES
OF THE GENUS CALLOPHRYS
J. W. TILDEN
San Jose State College, Sort Jose, California
INTRODUCTION
EXAMINATION OF CERTAIN SPECIMENS of Callophrys from mon-
tane habitats in California and Oregon indicate that these specimens
belong to a cluster distinct from others so far described, and not fitt-
ing well with any known description of North American Callophrys.
The purpose of this paper is to discuss certain characteristics of North
American Callophrys, and to provide a name for the previously un-
recognized series of populations. This name is needed for two forth-
coming faunistic papers.
TAXONOMIC HISTORY OF CALLOPHRYS
Linnaeus (1758) described P (apilio) P {lebeji) rubi as follows:
‘Rubi. 154. P. P. alls dentato-subcaudatis; supra fuscis, subtus viri-
dibus.”
Billberg (1820) proposes the genus Callophrys with three groups,
based on three tails, two tails and one tail. In the one-tailed group only
rubi is listed.
Scudder (1875), selected rubi as the type of Callophrys as follows:
'T820, Billb., Enum. Ins. 80: Vulcanus, rubi, and a Ms. species. Rubi
may be taken as the type.”
Until recently Callophrys has been limited to those few hairstreaks
that matched rubi in every respect. The palaearctic names, besides rubi
itself, include borealis KruL, polaris Moschl., fervida Stgr., sibirica
Ruhl and suaveola Stgr., omitting abberations that have been named.
All of these have been considered both as full species and as subspecies
or variants of rubi. Of these only rubi, as type of genus, will be men-
tioned further, "f
Ziegler (I960), combined the groups placed under Mitoura Scud.,
Incisalia Scud., Sandia Clench & Ehrlich, and Callophrys Billb., under
the oldest name, Callophrys, as subgenera. This move had been pre-
sented in abbreviated form previously by Ehrlich and Clench (I960)
in the paper describing Sandia mcfarlandi. Clench, in Ehrlich and
Ehrlich (1961) used a similar arrangement but several genera are
281
282
TILDEN
/. Res. Lepid.
proposed for groups of hairstreaks outside the scope of this paper,
which is concerned only with the members of the presently considered
subgenus Callophrys as limited by Scudder’s designation of rubi as the
type.
THE AMERICAN SPECIES OF THE SUBGENUS CALLOPHRYS
All species of the subgenus Callophrys resemble one another very
closely, and can be separated with any degree of assurance only by
considerable effort. There always remain specimens the position of
which is a matter of opinion in the light of present knowledge. There
are two revisions of the group. Barnes and McDunnough (1923),
grouped the species in a manner superior to previous treatments and
described three subspecies. Clench (1944) revised the genus (as it
was then considered) proposing one subspecies.
At present at least ten names are involved in the North American
Callophrys (s. str.), and these have been regarded at one time or
another as representing six distinct species. Characters that have been
used to distinguish the various species have been color markings and
wing shape. The male terminalia are exceedingly similar in all the
named entities. This is not to say that differences may not exist, but
so far they are not apparent. The present author has also dissected the
female genitalia of the North American species and the European
rubi. These studies have not resulted in any conclusions in time for
the present paper.
An attempt has been made to examine all parts of the external
surface of the body. Small differences may be detected in the palpi,
antennae, facial hairs or tuft, and in wing shape, stigma and color
markings. The results of these comparisons are given in Table 1.
It seems pertinent to include here a discussion of the problem of
the identity of dumetorum Bdv. Boisduval ( 1852) writes (free tran-
slation from the original French) that "this Theda resembles our
rubi in every respect and is most likely only a local variety of that
species.” Close examination of rubi in comparison to specimens usually
considered to represent dumetorum Bdv. shows a number of minor
differences, but without doubt the two are closely related. Certain
problems arise in deciding what would properly be considered as
dumetorum Bdv. Clench (1955, op. cit, p. 220) gives a description
of what he at that time regarded as dumetorum. The description
suggests that he is describing the insect for which a new name will be
proposed in this paper.
The Boisduval description could fit any of several named segre-
gates. The figure by Oberthur (1913) shows a gray insect with over-
scaling of a cast of green not usual in the subgenus, and with a rather
more complete macular band than one expects in insects that pass
for dumetorum in collections. W. D. Field, of the Smithsonian In-
(4):28i-}00, 196J
NORTH AMERICAN SPECIES
283
sitution, has kindly given me a detailed comparison of the insect
figured by Oberthur, and which is usually regarded as the type of
dumetomm. Whether or not this specimen is actually the one on
which Boisduval based his description is not certain; this does not
seem to be stated explicitly. Field- informs me that Oberthur’s figure
is a faithful representation of the supposed type specimen. This speci-
men is a gray female with the fringes white or pale-tipped with a con-
spicuous macular band and no antennae.
This female specimen, by virtue of being illustrated by Oberthur,
may need to be regarded as a lectotype, the illustration being one
method of designation, as was suggested by Field (in litt), A possi-
bility exists that it may indeed be the specimen described by Boisduval,
but this point can not be settled in light of present knowledge.
Neither Boisduval’s description nor Oberthur’s illustration seem
to fit exactly any presently known population of Callophrys. Either
dumetomm was described from unusual specimen, or from some
since unlocated population, or (most likely) neither the illustration
nor the description are completely recognizable in relation to some
well-known population. Of the specimens compared with the type,
there seem to be fewer discrepancies with the lowland population
most commonly considered to be dumetomm, except that the type is
a gray female while the females of the lowland population tend to
be brown or fulvous. However, gray females are not entirely un-
known. The macular band is more than usually well-developed in
the "type,” but the band is variable in all species, and specimens
from the foothills of the Sierra Nevada are known with complete
macular band. The choice here made is to continue to use the name
dumetomm for the cismontane lowland population of California, while
retaining the option of changing this opinion should future findings
make such a change desirable.
The exact locality where Lorquin collected his original specimen,
subsequently named dumetomm by Boisduval, is partially in doubt,
as is frequently the case with the Lorquin-Boisduval specimens.
Species described in 1852 were from Lorquin’s earlier collecting. In
some of the descriptions there are phrases such as "Mountains of the
Juba,” suggesting that the specimens were taken near the mining
operations along the Yuba River. A reading of Lorquin’s adventures
in California brings the same conclusion; that he was in the Sierran
mining region prior to 1852. This is presumptive evidence, but
certainly better than no inference at all. Specimens of Callophrys from
the Sierra Nevada foothill localities relate to the populations from
coastal and southern California rather than to those of the high Sierra
Nevada. Here again the evidence favors use of the name dumetomm
for the low elevation insects.
Should incontrovertible evidence subsequently be found, that this
284
TILDEN
/. Res. Lepid.
Character
dumetorum auct.
apama
viridis
wing fringes
fuscous basally;
pale tipped; not
clear white
dark fuscous basally;
pale tipped, dark
scales mixed.
fuscous; the tips
mixed pale and
fuscous scales
fore wing apex
obtuse-angled
obtuse-angled
obtuse-angled
forewing, outer
margin
oval in female;
straighter in male
and indented at Cu2
oval in female;
male slightly pro-
duced between Cu]^
and
curved to or M ;
then nearly straight;
tornus slightly
incurved
hindwing, outer
margin
crenate in spaces
Cuj^, Cu2 and 2nd A.
(indentations shallow)
small crenation in
space Cu2; a deeper
one in space 2nd A
hind wing appearing
more than usually
quadrate; crenations
very slight
color of fore-
wing costa
below
fulvous, more marked
in female
pale brown to dull
fulvous; concolorous
with other brown
areas of wing
rich fulvous in both
sexes
forewing below
invaded by gray or
fulvous from anal
margin to at least
vein M]^
anal margin gray;
disc invaded by rich
fulvous at least to
vein M]^
gray confined to anal
cells only
vein-tips of
hind wing below
vein tips and marginal
line usually rusty-
brown scaled
narrow but complete
marginal line, out-
brown, mesially
black, inwardly
white; vein tips dark
brown scales at vein-
tips absent or nearly
so
Macular band
(lower surfaces
of wings)
usually reduced to
3 spots; seldom com-
plete; macules in-
wardly brown, then
white
hindwing: complete,
spots in Cuj^ 6i Cu2
displaced out. Tri-
colored, 4-5 spots
forewing
complete on both wings
(though narrow) to re-
duced; mesial brown
scaling reduced
shade of green
below
grass green to
golden green
rather dull green,
brightened by ad-
mixture of fulvous
scales
deep, frequently
bluish, green.
scaling of
labial palpi
black above, mixed
black 6c white below;
may have few green
scales near base
hairs and scaling
smooth; black above
and below; white
scales laterally
grizzled black and'
white throughout;
third segment very
pointed; scaling
sparse, green scales
basally
facial hairs
(facial tuft)
erect or slightly
proclinate; thicker
laterally; black with
green scales at bases
hairs markedly pro-
clinate, thicker
laterally; basal
green scaling
erect or slightly
proclinate; thicker
laterally; black or
gray; green scales
at base
forewing
stigma of male
small, ovate, usually
gray and lighter than
ground color
ovate to oval;
gray, slightly
lighter than ground
small, ovate, dis-
tinctly paler than
ground color, or
concolorous
antennal annuli
(white rings)
usually 15 (seldom
16)
usually 15 (seldom
16)
14-17 (av. 16 in 34
specimens) ; antennae
pale above (unique)
general facies as
seen from above
male uniform fuscous
with pale- tipped
fringes; female brown
with fulvous discs
and pale- tipped fringes
male uniform dark
fuscous; female
same with very
large fulvous discal
areas
gray
insect with nearly
concolorous fringes;
females usually gray
(very seldom partly
brown on disc)
TABLE 1
CHARACTER COMPARISON OF NORTH AMERICAN CALLOPHRYS (S.STR.)
(4);28i-300, 1963
NORTH AMERICAN SPECIES
285
affinis
sheridani
corns tockl
lemberti n. sp.
basally fuscous;
pale- tipped to
white tipped
bases darker than
wing; tips mixed
white & fuscous
bases darker than
wing; tips white ,
contrasting
basal scales mixed )
brown and gray; tips
snow white (usually);
approx, right-
angled, tip rounded
acute, the forewing
trlgonate
rectangular; effect of
short sharp tip
pointed, nearly as in
sheridani!
curved to ,
thence nearly
straight 'to tornus
alight curve to
or M2, then straight
to tornus or
slightly concave
quite evenly curved,
the curvature
slight
curved to M3 ^
slightly indented
between Cuj^ & Cu2
quadrate, nearly
rounded; crenations
slight or obsolete
rounded; crenations
scarcely visible
longest between M3
& Cu^; crenations
evident but minute
rounded, crenations
slight, between CU2
& 2nd A one is
evident
gray to clay-
colored, not
contrasting
blackish; darker
than rest of wing
dark in most males;
concolorous to pale
fulvous in females
narrowly brown or
fulvous; contrasting
gray from anal
margin to Cu ;
general wing
surface green
dark gray to
vein Cu2
gray area extensive,
to M2 (one female) on
to across wing to
costa (one male)
gray confined to anal
cells; even here some
green overscaling
slight brown at
each vein- tip;
terminal line
not differentiated
spots at vein tips
not evident; term-
inal line black
dark vein tips not
evident; terminal
line black, narrowly
white Inwardly
usually slight dark
points at vein tips
on both pairs of
wings; approaches
checkering
obsolete, or one
spot in cell Cu2
complete; straight;
white; edged within
and without by black
band complete but of
separate spots; bowed
out at cell Cu2;mac-
ules white, black
inwardly .
^ complete band to
obsolete of discrete
white spots black
inwardly, narrow;
obscure on forewing
uniform pale
yellow- green
deep dark green scales
mixed with dark gray
scales about equally
deep dark green, the
veins slightly con-
trasting dark
green scaling thin,
luniform, bright pale
‘green, the undercolor
showing through
palpi slender
and thin scaled;
mixed black and
gray all over
dark above; scaling
below dense; mixed
black and light
gray (effect dark)
palpi slender,
pointed; scaling
sparse; dark above,
mixed black & white
below
palpi slightly darker
above but mixed white
and black scales
throughout; third
joint darker
facial hairs
light gray,
sparse, procumbent
dark, dense, slight-
ly proclinate;
underscaling pris-
matic, not green
tuft dense, coarse,
b^ack, proclinate;
subscaling covered
or obscure
hairs sparse, fine
dark, sub-erect;
green sub-scaling
prominent
dark to black,
sharply con-
trasting
stigma usually
nearly concolorous
with wing
small, slightly pale
to concolorous and
scarcely discernible
stigma small, sub-
triangular, con-
colorous to pale;
seldom dark
17
av. 17 (16-18)
15-16
16 (17 on one
specimen)
fulvous to bright
rufous with dark ter-
minal line & pale
fringes
dark gray with black
terminal line and
white fringes; sexes
alike in color
gray insect with
dark terminal line
and white fringes;
sexes concolorous
mouse gray smooth-
scaled insect with
white fringes; females
concolorous or dull
brown (fuscous)
TABLE 1 (continued)
286
TILDEN
/. Res, Lepid.
rubi
Incisalia augustinus
Mitoura siva
Sandia mcfarlandi
basally dark; tips
with a few pale
hairs
mixed brown & gray
basally; tips sordid
gray; dark at vein-
tips (checkered!
forewings fuscous;
hindwings pale- tipped
basally brown and
fuscous mixed; tips
white
tip quadratCji bluntly
pointed
tip rectangular or
nearly acute
rather acute
obtuse
rounded to thence
nearly straight to
tornus male indented
at Cu2
rather evenly curved
but slightly flatter
before tornus
quite evenly rounded
slightly flattened
before tornus
rounded
two well-marked
crenations at cells
Cu2 and 2nd A.
at least a suggestion
of a crenation between
each pair of veins
tailed at Cuj^ & Cu2;
not crenate
tornal crenation
evident, slight
broadly fulvous
in female; narrowly
so or gray in male
concolorous with wing
slightly fulvous, not
greatly contrasting
lemon yellow to
apricot
gray confined to anal
cells
no contrast
fulvous invasion from
anal margin almost to
costa
yellow to apricot
shade across entire
wing
vein- tips brown;
terminal line warmi
brown in female,
less so in male
terminal line dark
brown to black; veins
not contrasting
complex pattern of
black, white and
rufous overscaling
terminal line black,
invading vein tips
of small separated
narrow white spots;
nearly complete to
obsolete
complete band of
small round dark
spots; dark basal
shade
complete, irregular
from inward out of
bands of brown,
black & white
narrow, complete,
white flanked each
side with black
grass green-
suggests dumetorum
ground color
pale vinaceous brown
smooth pale green
scaling where other
patterns are not
evident
basically a luminous
yellow- green
mostly dark with
white scales on
sides; green scales
usually evident
mixed black & white
scales; darker above
black above and at
tips; basally, white
overscaling
dark above and at
tips; white scaling
below and laterally;
rather long
sparse, dark, mostly
lateral; median
green sub-scaling
prominent
dense dark rich
brown, procumbent;
sub-scaling covered
very sparse, pro-
cumbent; iridescent
underscaling very
visible
hairs fine, short
& erect; mixed
black & white;
subscaling dark
elongate-oval ;
androconia rather
rough
very long oval,
androconia small;
black to pale
elongate-oval ;
dark to pale
elongate, about
3 times as long
as wide
16
18
1 7 , narrow and
clear-cut
15-16; antennae very
short, each segment
gray with nearly
concolorous fringes
(male) ; dark brown
with concolorous
fringes (female)
dark brown with
checkered fringes
(male) ; female
lighter richer brown
dark brown Insect
with fulvous on discs
and two dark dots
near hind wing tornus;
female more fulvous
brown insect with
fulvous discs &
white fringes
TABLE 1 (continued)
I (4) :28i.300, 196}
NORTH AMERICAN SPECIES
287
position is untenable and that the population herinafter described as
unrecognized is actually true dumetorum, the populations now con-
considered as dumetorum and its variants must then become known
as perplexa B. & McD., as the oldest available name definitely assigned
to this coastal insect.
The notations in Table 1 are of necessity short. In some cases
amplification is given in the species discussions. The characters used
in Table 1 vary in the samples from the different populations, but
the degrees of quality given are averages. In all cases the samples were
adequate (twenty-five or more specimens) to large (fifty to one
hundred specimens) except for af finis and comstocki. Of the latter,
only four specimens were available for close study, and some fifteen
specimens were examined in all.
For comparison, one species each of the subgenera Incisalia, Mitoura
and Sandia (which contains a single species) are included and com-
parison will show that most of the characters used are specific
rather than generic or subgeneric. The subgenus Narnia was excluded
from lack of material, and the subgenus Cyanophrys because it enters
our fauna only along the extreme southern border.
Clench (in Ehrlich & Ehrlich) defines Callophrys (s. lat.) and
its subgenera primarily on genitalic characters. The subgenus Callophrys
includes species with valvae not capped; the cornuti of the aedeagus
slender, not spatulate; the scent-pad (stigma) well-developed, and
the labial palpi about IV2 times as long as the vertical eye diameter.
(This last character is shared with the other subgenera except for
sandia, the palpi of which are about twice as long as the vertical
diameter of the eye). The scales of the stigma are entire and with
rounded ends and the hind wing is not tailed.
In addition may be mentioned the tornal "tab” of the hind wings,
shared with Incisalia; the reduction of the hind wing marginal crenula-
tions, restricted usually to the last three cells (much more extensive
in Incisalia); the usually even green overscaling of the inferior sur-
faces, typical of the subgenus, and the complete lack of the thecla
spot. The markings below are restricted to the submarginal band
(macular band as frequently stated), which is often reduced or even
absent, even in a single species. There is no other evident ornamenta-
tion.
In the following analyses of the species, only the original citation
is given.
Callophrys rubi ( L. )
Syst. Nat. 10th Ed. 1:483, No. 154:1758
Palaearctic. Range Europe and Asia. By some considered the only
valid palaearctic species. Closely resembles dumetorum as considered
here, but the green darker, the female less broadly fulvous, the wing
288
TILDHN
/. Res. Lepid.
fringes less contrasting, the stigma more elongate, and the under sur-
face of the forewing not greatly invaded by gray or brown. In all
specimens examined, the terminal line of the wings is much more
evident.
Gallop hrys dumetomm (Bdv.)
Ann. Ent. Soc. France (2) 10:291:1852
A fuscous (male) or usually broadly fulvous (female) insect with
pale-tipped but not white fringes; hind wing with three crenations,
terminal line not strongly contrasting. Green below warm grading to
yellowish green in southern specimens, nearly distinctive. Forewing
deeply invaded by gray (male) or tan (some females) at least to vein
Ml and frequently (subspecies perplexa) to forewing costa. Veins
of secondaries below dark tipped, the border usually clouded with
brown scales. Whether or not this species is correctly identified as
dumetomm, it is one of the distinctive entities of the subgenus. The
invasion of the forewing by gray or brown seems diagnostic in its
range. The subspecies perplexa B. & McD. (1923) is more yellowish-
green below, the band reduced or obsolete (usually) and with the fore-
wing invasion frequently extending to the costa. However, reduction
of the macular band occurs in all populations of dumetomm (as well
as in nearly all other populations of Callophrys) and this character
must be used with caution. The macules of the band are bicolored,
brown inwardly, outwardly white.
Callophrys apama ( Edw. )
Papilio 2:137:1882
A very dark fuscous (male) or fuscous with broad fulvous discs
(female) insect with fringes slightly pale-tipped, not notably contrast-
ing. Green below rather dull but mixed with fulvous scales, giving
a superficial appearance of being lighter than is really the case. Macular
band (nominate apama) complete, the macules in spaces Cui and
Cug displaced outwardly, the macules of the band tricolored, inwardly
brown, mesially black and distally white. Facial tuft peculiar, sparse,
rather light in color and the hairs markedly proclinate in all of the
specimens examined. The name homoperplexa B. & McD. (op. cit., p.
68) was given to specimens such as those from Colorado, in which the
macular band tends to become obsolete. The shade of green and the
other characters remain similar.
Callophrys viridis ( Edw. )
Though usually considered a synonym of dumetorum, viridis appears
to be as distinct as most of the species, and was ressurected from
synonomy by Clench (1944). It is a dark gray insect in both sexes
when fresh (old specimens fade to a lighter gray), the fringes pale-
(4):28i-500, 1963
NORTH AMERICAN SPECIES
289
Fig. 1. Callophrys. Upper side to left; lower side to right. Callophrys
mbi L. Near Hannover, Germany, 12:IV.52. G. Hesselbarth.
Callophrys dumetorum Bdv. El Portal, Mariposa Co., Calif. 16. IV. 61, ca. 1200’,
J. W. Tilden.
Callophrys viridis Edw. Twin Peaks, San Francisco, Calif. 16. III. 57, J. W.
Tilden.
Callophrys affinis Edw. Dry Canyon, Salt Lake County, Utah 14.VII.49, J. C.
Downey.
Photography by Jean Norton
290
TILDEN
J. Res. Lepid.
tipped and usually contrasting, particularly on the hind wings. Occasion-
al females in a long series are brown, very seldom fulvous, and nor-
mally females cannot be separated from males on color alone. The
outlines of the wings are more quadrate than in other members of
the subgenus; the crenations of the hind wings are discernible but
small. The forewing below has the gray limited to the anal margin,
not invading the disc. The green below is dark and frequently bluish,
quite distinctive. The antennae are pale to whitish above, seemingly an
unique character. The range in California is rather narrowly limited
to the coast, from Santa Cruz County northward. Its range beyond
California needs to be clarified. It flies very early in the year, late
February to early April, and seldom leaves the vicinity of Eriogonum
(usually E. latifolium Sm.) except to visit nearby flowers. It is partial
to flowers of Umbelliferae. Its flight is low and easily overlooked. In
spite of reports to the contrary, viridis seems to be the only species of
the subgenus found in the immediate environs of San Francisco, and
is also common in similar habitats in Marin County.
Gallop hrys af finis ( Edw. )
Proc. Acad. Nat. Sci. Phila., 223:1862
A fulvous to bright rufous insect, with dark terminal line and
usually dark contrasting stigma, the fringes pale-tipped to white,
usually very contrasting. Below, uniformly pale yellow-green (nom-
inate af finis), the macular band obsolete or represented by one or two
minute macules only. This describes nominate material from Utah.
North and west, specimens associated with this species are less fulvous
above, more bluish-green below. W ashingtonia Clench (1944) is based
upon such specimens from Brewster, Washington.
Callophrys sheridanii ( Edw. )
Eield and Eorest 3:48:1877
Whether this name is to be attributed to Edwards or to Carpenter
may be a matter of opinion, A rather short badly written article with
several misspellings and typographical errors, by Carpenter, states that
Edwards is describing the species. It reads: 'Theda sheridonii (sic),
new species, is named in honor of Lieut. Gen. P. H. Sheridan, U. S.
Army, by W. H. Edwards, Coalberg, West Virginia, at the request
of W. L. Carpenter, U. S. Army. Size and form ...” Inasmuch
as it is expressly stated that Edwards is writing the description and
that it is at the request of Carpenter, the position is taken here that
Edwards is the author of the name. However, the alternate opinion
has also been expressed. In any case the present spelling is an emenda-
tion of an evident lapsus calami.
(4)s28i-30o, 1963
NORTH AMERICAN SPECIE^
291
Fig. 2. Callophrys. Upper side to left; lower side to right,
Callophrys sheridanii Edw. Flagstaff Mt., Boulder Co., Colo. 17.IV.52, Don
Eff.
Callophrys apama Edw, Strayhorse, Greenlee County, Ariz., 7.VII.58, 7800
ft., J. W. Tilden.
Callophrys comstocki Henne Providence Mts., San Bernardino Co., Calif. 2. IV.
50, Ray Hulbirt.
Callophrys lemberti Tilden, n. sp. West above Tioga Pass, Yosemite National
Park, Calif. 8.VII.58 ca. 10500 ft., O. Shields.
Photography by Jean Norton
292
TILDEN
/. Res. Lepid.
Sheridanii is the most divergent species in an otherwise closely re-
lated group. It is dark gray above in both sexes, with a black terminal
line and white-tipped contrasting fringes. The stigma of the male
is small, usually concolorous and not immediately evident. Below,
the green is very dark and mixed almost evenly with black scales. The
forewing is more acute than in the other species and the secondaries
appear smaller and more evenly rounded. The facial tuft is unusually
dense, erect and black, and most specimens show eighteen annuli in
the antennae. The macular band is only slightly arcuate and is not
usually broken into discrete macules. It is white mesially, faced on both
edges with black. The name neoperplexa B. & McD. (Contrib. 5:671;-
1923) was applied to specimens from Utah in which the macular
band tends to become reduced or obsolete. The western range of
neoperplexa remains to be established, but it appears to extend into
eastern Oregon and eastern Washington.
Callopbrys cornstocki Henne
Bull. So. Calif. Acad. Sci. 39:71:1940
This species was described from the desert region of San Bernardino
County, Calif. (Providence Mountains) and specimens are relatively
scarce in collections. The precarious climate results in good populations
of adults only in favorable years. Comstocki has been considered either
a distinct species or a subspecies of dumetornm. Examination of the
short series available to me for study, while not conclusive, indicates
that co7nstocki is separable from other named segregates of the sub-
genus by characters at least equal to those defining most of the species.
It is a gray insect in both sexes, the terminal line dark, the fringes
white tipped and contrasting at least on the secondaries. The green
below is dark, the veins slightly darker than the background color.
The stigma is small, scarcely discernible and the facial tuft is dense,
coarse, black and proclinate, almost concealing the underscaling. The
vein tips are not dark on the hind wing below, but the terminal line
there is black, nearly complete and inwardly bordered narrowly with
white. The forewing is deeply invaded by gray, a character (almost
the only one) it shares wiuh dimietoriim. The macular band is com-
plete but of discrete macules, inwardly black, outwardly white, and
the spot in cell Cu2 is displaced outwardly.
Callopbrys lemberti Tilden, n. sp.
A mouse-gray smoothly scaled species with pale or concolorous
stigma and contrasting white-tipped fringes. Sexes similar, or females
dull brown. Green of lower surfaces pale, the scaling thin, the ground
color showing through. Macular band complete to obsolete, unusually
narrow, of discrete macules.
Holotype male: Costa of forewing 13 mm.; costa upcurved to basal
(4):28r-300, 1963
NORTH AMERICAN SPECIES
293
Fig. 3. Map showing distribution of CaUophrys apama (°) and C. sheri-
danii ( • ) .
294
TILDEN
/. Res. Lepid.
third of cell, then straight to end of R2 + 3, curved down thence to
apex; outer margin of forewing curved to M3, thence nearly straight
to 2nd A except for a slight indentation between Cui and Cus; tornus
diagonal, anal margin nearly straight. Hindwing: Rs=2W A; Mi =
Cui, these the longest veins in this wing; two slight marginal crenations
between Cui and Cu2 and between Cuo and 2nd A; white annuli of
antennae 16 as seen from lateral view; palpi dark above, grizzled
black and white on sides and below, rather more blunt than usual
in the subgenus but of no significance, since other specimens of the
type series have palpi more pointed than in the holotype; facial area
with gray, slightly proclinate hairs, the vestiture more dense laterally
and with few irridescent green subscales showing through; body dark
above, pale below; the legs annulated dark gray and white.
Upper wing surfaces gray (nearly mouse gray), the veins very
slightly darker but not greatly contrasting; stigma nearly concolorous
with wing; fringes of forewing concolorous at base, white at tips;
fringes of hind wing concolorous gray mixed with golden brown scales
at base, snow-white and contrasting at tips; tornal table downturned
and dark.
Lower wing surfaces with smooth, pale, slightly yellowish green
(nearly apple green) overscaling, which is thin, the gray ground
color showing through between the individual scales, costa of fore-
wing pale brown, moderately constrasting; anal area of forewing
gray from margin to Cu2, the gray not invading the disc; forewing
with a slight suggestion of ' a macular band with macules in cells
M3, Cui and Cu2; fringes as on upper surface.
Secondaries wdth green overscaling over entire surfaces; macular
band nearly complete but narrow, the macules narrowly black in-
wardly, outwardly and more widely (about Vd) white; the macule in
cell ^rd A is a short dash; that in cell 2nd A directed diagonally toward
wing base; that in cell Cui. also diagonal but displaced towards wing
margin by about one-half its own length; macule in cell Cui lacking
on right wing, indicated by four or five white scales on left wing;
that in cell M3 very faint, dull white and narrow; that in cell M2
faintly indicated by a lack of green overscaling only; no macule in
cell Mi; macule in cell Rs small but distinct; fringes nearly as
on upper surface except for clusters of dark scales at vein tips, sug-
gesting incomplete checkering; the tornal tab centrally black, narrow-
ly faced on each side with white hairs; hairs of vannal margin gray
at end of macular band, thence nearly white to base.
Allotype female: Forewing costa 13.5; wing shape and proportions
essentially as in holotype male; white annuli of antennae 16 (an
incomplete 17th on base of club); facial area with hairs sparse (as
is frequent in female Callophrys) ; Body and legs as in holotype male.
Upper surfaces dull gray-brown, darker and very slightly more
(4):28i-3oo, 1963
NORTH AMERICAN SPECIES
295
Fig. 4. Map showing distribution of Callophrys viridis (1), lemberti (2),
and comstocki (3).
296
TILDEN
/. Res. Lepid.
brownish than holotype male. Costa above with warm brown scales,
some of which invade the wing nearly to vein Sc; fringes of fore-
wings dark gray at base, white at tips, of hind wings brownish at base
except for dark gray patches at vein tips, the fringe tips white; tab
down-turned and dark, the fringes in the tornal area invaded by
gray, the tips less contrasting.
Green overscaling of lower surfaces as in the male; forewing
costa warm brown (nearly cinnamon); macular band indicated by
two indistinct macules, in cells M3 and Cu,; fringes light gray at
base, with darkenings at vein tips; fringe tips white; gray anal margin
free of green overscaling not quite to Cus; macular band of hind
wings very narrow, the spots small and white with the inward black
scaling scarcely more than suggested, the band slightly bowed out
but the macules not noticeably displaced; macules in cells M2 and M3
obsolete; fringes pale gray, distinctly dark at vein ends, appearing
checkered; fringe tips white; a faint subterminal row of paler green
scales before the fringes; tab and vannal margin as in holotype male.
Type material: Holotype male. West above Tioga Pass, Yosemite
National Park, Calif., 9:VII.62, leg. Oakley Shields; allotype female,
same locality, 10.VII.58, leg. Oakley Shields; seven designated para-
types as follows: 1 male 8.VII.58 (Shields); 2 males, + 1 $ 19. VII. 52
(Tilden); I male 25.VI.6l, -j- 1 9 (Dirks); 1 male, 25.VI.62
(Dirks): All paratypes from same locality. Type locality: West above
Tioga Pass, about 1 mile, where the Gaylor Lakes Trail reaches its
highest point before dropping down to Gaylor Basin, thence southerly
along the ridge to rock oucrops, about two to three hundred yards.
Type material distributed as follows: Holotype male and allotype
female in the collections of the California Academy of Sciences; one
male paratype in the collection of Oakley Shields, La Mesa, Calif.,
one male paratype in the collection of the Los Angeles County
Museum; one female paratype in the collection of the Carnegie
Museum, Pittsburgh, Penn.; one male paratype in the collections
of the National Museum, Washington, D. C. The males are retained
by the author because they have been dissected to examine the geni-
talia. Certain other specimens from the type locality are at hand but
are in too poor condition to form paratypes.
Variation in the type series: In flown specimens the green appears
very slightly darker; the macular band is never more complete or
conspicuous than in the types. In three paratypes the macules of the
band are smaller and in one specimen the band is very faint. One
male paratype has the stigma very pale; in one other the stigma is
slightly darker than the ground color.
Recognition characters for Callophrys lemherti, n. sp. are the smooth
gray upper surfaces, nearly similar in both sexes; the very contrasting
white-tipped fringes, particularly on the hind wings; the very narrow
NORTH AMERICAN SPECIES
297
I (4) ;28i-3oo,
1963
Fig. 5. Map showing distribution of Cdlophrys af finis.
298
TILDEN
J. Res. Lepid.
but frequently nearly complete macular band; the sub-checkered
fringes of the hind wings below. It may be separated at a glance from
dumetorum by the smooth gray upper surfaces and white fringes,
and by the restriction of the gray of the lower surfaces of the
hind wing to the anal margin. From viridis it may be separated by
smaller size, more trigonate forewing and the pale green scaling below.
From comstocki is may be known by the lack of contrast between
ground color and the wing veins, especially on the lower surfaces of
the hind wings, and by the more regular and narrower macular
band and the contrasting brown coloration of the forewing costa. It
bears no confusing resemblance to the remaining species. Specimens
of lemherti, n. sp., collected long ago, tend to fade, the females parti-
cularly showing a sordid gray-brown above. The tendency of Callo-
phrys specimens to fade is general.
Specimens from other localities than the type locality referred
to lemberti, n. sp., are: I male, 1 female, west slope Mt. Dana, Yose-
mite National Park (Tilden); 1 male, I female, Warren Creek, Mono
County, Calif. (Shields); 1 female. Mammoth Crest, Mono Co., Calif.
( J. Powell ) ; 2 males 1 female. Chipmunk Flat, Tuolumne County,
Calif. (J. Powell); 2 males 1 female, the knobs just north of Sonora
Pass, Stanislaus County, Calif. ( Shields ) ; 1 male 6 females, Ebbetts
Pass, Alpine County, Calif. (J. Powell); 1 male I female, Leviathan
Peak, Alpine County, Calif. (J. Powell); 1 male. Echo Lakes Area,
Eldorado County, Calif. (Dirks); another male from Echo Lakes,
without collector label; 1 female, Mt. Tallac, Eldorado County, Calif.
(F. X. Williams, 1909); 1 male. Tamarack Lake, Eldorado County,
Calif, (no collector label); 1 male. Crater Lake National Park, Kla-
math County, Oregon (D. Huntzinger); 1 male, west slope Mt.
Thielson, Klamath County, Oregon (Shields), 26 specimens in all
extending from the Central Sierra Nevada to nearly central Oregon.
In addition there are several apparently conspecific specimens too
worn to use for reference.
The species is named in honor of an early collector in the Tuolumne
Meadows and Tioga Pass sections of what is now Yosemite National
Park.
Thanks are due the following individuals and institutions for the
loan of material: Ernst Dornfeld, Corvallis, Oregon; David Huntzinger,
Mt. Timpanogos Cave National Monument, Utah; C. Don MacNeill,
California Academy of Sciences, San Francisco, Calif.; E. J. New-
comer, Yakima, Washington; Jerry Powell, California Insect Survey,
University of California, Berkeley, Calif.; Oakley Shields, La Mesa,
Calif., and Fred Thorne, El Cajon, Calif.
Special thanks are due Paddy McHenry, Burbank, Calif., for pro-
viding certain references not otherwise available.
C4)s2Si-30O, 1963
NORTH AMERICAN SPECIES
Fig. 6. Map showing distribution of Callophrys dumetomm.
300
TILDEM
/. Res. Lepid.
SUMMARY
The species of Callophrys Billberg, s. str., are compared by use of
certain minor characters mostly previously unused. An unrecognized
species, C. lemherti, n. sp., is described. Specimens other than those
of the type series are referred to this new name and the range as
now known is given, being from the central Sierra Nevada of Cali-
fornia to Mt. Thielson, Oregon.
REFERENCES
BARNES, W., & J. McDUNNOUGH 1923. Contrih. Nat. Hist. Lepid. North
America. 5:64.
BILLBERG, C. J. 1820. Enumeratio insectorum in Museo Billberg, p. 80.
BOISDUVAL, JEAN A. 1852. Lepidopteres de California. Ann. Soc. Ent.
France, (2) 10:291.
CLENCH, H. 1944. Notes on lycaenid butterflies, a. The genus Callophrys
in North America. Bull. Mus. Comp. Uoo. 94:217-229.
CLENCH, H., in EHRLICH, P. R., & ANNE H. EHRLICH. 1961. How to
know the Butterflies, 200-211.
EHRLICH, P. R. & H. CLENCH. I960. A new subgenus and species of
Callophrys (s. 1.) from the southwestern United States. Ent. News
71:137-141.
EDWARDS, W. H. 1862. Proc. Acad. Nat. Sci., Philadelphia, p. 223.
. (in CARPENTER, W. L.) 1877. Field & Forest, 3:48.
. 1882. Papilio 2:137.
HENNE, C. 1940. Bull. So. Calif. Acad. Sci., 39:71.
LINNEAEUS, C. 1758. Systema Naturae, 10th Ed., 1.483.
OBERTHUR, C. 1913. Etudes Lep. Comp., Ease. IX (Partie Ire) PI. CCXXX-
VI, fig- 1926.
SCUDDER, S. 1875. Proc. Am. Acad. Arts & Sci., Boston, 10:132, no. 202.
ZIEGLER, J. B. i960. Preliminary redefinition of North American Hair-
streak genera. J. Lep. Soc., 14:19-23.
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THE JOUI^NJAL OF KESIAR^OHJ
ONJ THE LEFIJOOFTERA
Volume 1 Number 4 May, 1963
IN THIS ISSUE
Techniques in the study of population structure in
Philotes sonorensis
Rudolf H, T. Mattoni and Marvin S. B. Seiger 237
Early stages of a southern California Geometrid moth,
Drepamdatrix hulsti huhti (Dyar) John Adams Comstock 245
The effectiveness of different isothiocyanates on attracting larvae
of Eteris rapae
William Hovanitz, Vincent C. S. Chang and Gerald Honch 249
The origin of a sympatric species in Colias through the aid of
natural hybridization William Hovanitz 261
A method for breeding Pieris napi and Pieris hryoniae
Bjorn Petersen 275
An analysis of the North American species of the genus
Callophrys J. W. Tilden 281
THE LEPIDOPTERA FOUNDATION 301
Notice 302
BIOGRAPHICAL SKETCHES
A number of these have been received but we need more. The space
where these could go is lost at the end of papers unless authors send in their
photos and sketches. We hope to start this feature in volume 2.
Established in 1 962
Edited by WILLIAM HOVANITZ
Volume 2
1963
Published at
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Volume 2
Number 1
July, 1963
YOSEMITE BUTTERFLIES: An ecological survey of the
butterflies of the Yosemite sector of the Sierra Nevada,
California ..... John S. Garth and J. W. Tilden 1
Volume 2 Number 2 September, 1963
Quantitative analysis of certain wing and genitalia characters
of Pieris in western North America
Vincent C.S. Chang 97
Biographical sketch — Vincent C.S. Chang 126
Genetic and environmental variation in Pieris brassicae
Brian O.C. Gardiner 127
Type localities of the Megathymidae H. A. Freeman 137
Biographical sketch — H. A. Freeman 142
The distribution of an endemic butterfly Lycaena hermes
Fred Thorne 143
Callophrys (Lycaenidae) from the Northwest
Harry K. Clench 151
Euphyes dukesi Bryant Mather I6I
The complete life history of Staphylus hayhursti
John Richard Heitzman 170
Volume 2
Number 3
November, 1963
Affinities and distribution of Antillean Ithomiidae
Richard M. Fox 173
Ovipositional preference tests with Pieris
William Hovanitz and Vincent C.S. Chang 185
Notes on the early stages of Drepanulatrix monicaria
(Guenee) ( Geometridae ) John Adams Comstock 201
Biographical sketch — John Adams Comstock 204
The origin of a sympatric species in Colias througli the aid
of natural hybridization William Hovanitz 199
Biographical sketch — James Wilson Tilden 224
A rubber stamp method for producing specimen labels
Phillip A. Adams 225
Generic or subgeneric names closely related to Argynnis
Paddy McHenry 229
THE LEPIDOPTERA FOUNDATION 240
Volume 2 Number 4 December, 1963
Evidence for lack of territoriality in two species of Hamadryas
Gary N. Ross 241
A synopsis of the west Indian Lycaenidae, with remarks on
their zoogeography Harry K. Clench 247
The synonymy, variability and biology of Lycaena nivalis
E. J. Newcomer 271
Comparison of the selective effect of two mustard oils and
their glucosides to Pieris larvae
W. Hovanitz and V.C.S. Chang 281
The Annaphila astrologa complex, with descriptions of three
new species Frank P. Sala 289
Old master painting Clark, Nelson of London, Ltd. 302
Dates of publication of volumes 1 and 2 302
Collecting of Annaphila spila with notes on the "crimson-
winged” group of the genus J. S. Buckett 303
Volume 2
Number 1
]ul^, 1963
THE JOUKNJAL @F RISEAKCHJ ©H
THE LEPIJDOFTEKA
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1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
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2(l):l-96
Journal of Research on the hepidoptera
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Copyright 1963
YOSEMITE BUTTERFLIES
An Ecological Survey of the Butterflies of the
Yosemite Sector of the Sierra Nevada, California
JOHN S. GARTH
Allan Hancock Foundation
University of Southern Calif orna
Los Angeles, California
and
J. W. TILDEN
San Jose State College
San Jose, California
July, 1963
2
GARTH AND TILDEN
/. Res. Lepid.
FOREWORD
Organized interpretation of the chief features of National Parks had its
beginning in Yosemite National Park in 1920. Suitable guide leaflets and
bulletins found a useful place in the educational program. At first printed
material covered wide fields of nature study. But with more visitors have
come many with specialized backgrounds and they seek help. Whereas, the
first bulletins attempted coverage of all insects found, we now come to one
dealing only with butterflies. The senior author graduated from the Yosemite
School of Field Natural History with the Class of 1933 and made his first
contribution to the subject by publishing a list of the Butterflies of the
Boundary Hill Research Reserve (Bull. Southern California Academy of
Sciences, 33, Sept.-Dee., 1934, pp, 13T135). He widened his studies to the
Butterflies of Yosemite National Park in the fifties and herewith is a paper that
emphasizes where each butterfly is found and what plant its larvae feed upon.
Colored plates give the amateur a chance to identify species found and the
text places zonal locations and ecology. Other national parks that have
profited by the senior author’s field work are Grand Canyon, Glacier, and
Grand Teton. Crater Lake and Lassen have profited similarly from the field
work of the junior author.- — Harold C. Bryant.
The original costs of this publication were underwritten by the
Lepidoptera Foundation and the Journal of Research on the Lepidoptera
and the separate issue is made available as a public service to visitors of
the Yosemite National Park and adjacent areas.
William Hovanitz, Founder, The Lepidoptera Foundation
Editor, The Journal of Research on the Lepidoptera
(c) Copyright 1963
Additional copies may be obtained by writing to THE LEPIDOPTERA
FOUNDATION, 1140 W. Orange Grove Avenue, Arcadia, California.
ALLAN HANCOCK FOUNDATION CONTRIBUTION Number 248.
2(i):i-$6, 1963
YOSEMITE BUTTERFLIES
3
CONTENTS
Foreword 2
Contents 3
Historical Note 3
Introduaion 5
The Lepidopterist’s Yosemite 6
The Surrounding National Forests 7
Yosemite Foot-Trails 8
Life-zones 9
Butterflies as Life-zone Indicators 9
Biotic Provinces .......................................................................13
Plant Communities 14
Food Plants ...........................................16
Some Plants on Which Yosemite Butterflies Feed as Larvae ..........17
The Geological Background ..........18
Acknowledgments ..............19
Names Used in This Paper ..19
Account of Species of the Yosemite Region 22
Literature Cited ........................68
Yosemite Butterfly Records ............................70
Appendix 1. Food Plant Records ....................................................71
Three Anomalous Situations ............................................................................77
Appendix 11. Yosemite Records ......................................................................78
Api^ndix III. Localities ..................................................................................92
Lepidoptera Foundation ....................................................................................96
HISTORICAL NOTE
The authors are indebted to Mrs. Mary V. Hood, Park Historian,
Yosemite National Park, for calling their attention to an early record
of butterfly collecting by John Muir, as recorded by W. F. Bade in
Life and Letters of John Muir, 1924:
Muir to Mrs. Ezra S. Carr, August 13, 1871. "I suppose you have seen
Mr. King, who kindly carried some [butter]flies for Mr. Edwards ... I collert-
ed most of them upon Mount Hoffman, but was so busy in assisting Reilly
that I could not do much in butterflies.” (Op. cit., Vol. I, p. 292).
Henry Edwards to John Muir, August 25, 1871. "In the small box that
you sent me [through Mrs. Carr] are four species new to my collection and two
of these ate new to science . . . All the specimens are rare, and are different
from those found in the Valley. The two new species are the bright crimson
copper one from Cathedral Peak, and one of the small bluish butterflies. There
is a pair of greenish yellow ones, very rare and interesting. The species was
described from a pair only which were taken by the Geological Survey at the
headwaters of the Tuolumne River, and strange to say, no others have turned
up until you found it now ... It is really very singular that the remove of a
few miles from the Yosemite should produce species so very different from
those of the Valley itself, and at the same time so charaaeristic in their forms.”
(Op. cit., pp. 263-264).
Of the butterflies mentioned, the bright crimson copper is probably
The Lustrous Copper, Lycaena cupreus (Edw.), while the greenish yel-
low ones are undoubtedly Behr’s Sulphur, Colias behrii Edw. It should
be noted, however, that the author is W. H. Edwards in each case.
Both species are illustrated on color plate III.
I^IITBTICS
r,i i; (I r't
4
GARTH AND TILDEN
/. Ris. Lepid.
Frontispiece. Boundary Ridge, el. 9,000 ft.. Research Reserve. Western
White Pine and sparse ground cover of herbs and grasses. Hudsonian life
zone elements weakly present. Rocky outcrop association. Papilio zelicaon
flies to the summit, while Euchloe creusa hyantis and E. ausonides coloradensis
keep to the leeward of the ridge.— -Neva Snell.
3(i):i-96, 196}
YOSEMITE BUTTERFLIES
5
INTRODUCTION
The Yosemite National Park embraces 1,189 square miles of the
most diversified territory to be found in the Sierra Nevada. From the
edge of the San Joaquin Valley at near sea level it extends to the crest
of the Sierran divide, culminating in Mt. Conness, Mt. Dana, and Mt.
Lyell, each over 13,000 feet in elevation. Between these extremes
occurs a wide range of climatic conditions, giving rise to a wealth of
plant and animal life scarcely to be duplicated elsewhere on the North
American continent. The vertebrate fauna has been thoroughly investi-
gated by Grinnell and Storer (1924), who recorded 97 species of
mammals, 231 of birds, and 34 of reptiles and amphibians in "Animal
Life in the Yosemite," their Yosemite sector extending beyond park
boundaries to include part of the San Joaquin Valley and of the Mono
Basin as well.
The larger host of invertebrates, and of insects in particular, have
long demanded similar investigation. The butterflies are an ideal group
with which to begin this study. They are diurnal, or daylight fliers, and
so lend themselves to easy observation, as distinguished from moths,
of which most are nocturnal or crepuscular in flight. They are also for
the most part conspicuous, and therefore likely to attract the attention
of even the casual park visitor. Again, their number as species found
in a given area roughly approximates the number of bird species, if
only those actually residing (i.e., nesting) in that region be enumer-
ated. Thus the butterflies constitute a natural group of insects small
enough to be encompassed by the amateur naturalist in a season, yet
sufficiently diversified to entice the professional entomologist to return
year after year to resolve the problems of habit and habitat that they
present.
Like the many other natural attractions of our National Parks, but-
terflies enjoy the protection of far-sighted legislation and are held in
trust for future generations as part of America s outdoor heritage.
Their unauthorized removal is contrary to park regulations; however,
the advanced researcher who can establish his connection with a public
museum or other scientific institution, such as a university, may qualify
for one of two types of collecting permits, and should consult the Park
Naturalist, with offices in the Yosemite Museum, regarding his eligi-
bility for this consideration.
6
GARTH AND TILDEN
/. Res. Lepid.
THE LEPIDOPTERIST’S YOSEMITE
The territory now included in Yosemite National Park has been
known to butterfly enthusiasts since the days of the gold rush of ’49.
The pioneer entomologist, Pierre Lorquin ( for whom Lorquin’s
Admiral is named), followed the mother lode to Placerville, and from
thence crossed the Sierra. His exact route is not known, but if he
crossed by Tioga Pass, el. 9,941 ft., it is quite possible that the "Great
Salt Lake" above which he discovered the Irene Fritillary (Argynnis
atlantis trene) was Mono Lake, and not the Great Salt Lake of Utah.
Indeed, had type localities been designated as carefully then as now,
we might find that many of our California butterflies were first ob-
served by a naturalist somewhere in the Yosemite region. The hermit
Lembert (whose name is memorialized in Lembert’s Dome) for many
years supplied the entomological world with specimens of Behr’s
Sulphur {Colias hehrii) from his hideaway in Tuolumne Meadows,
and it was not until the Tioga Road opened to tourist travel in 1915
that his secret source of this rarity was discovered.
The entire northern section of the park and much of the south-
eastern part is still traversable by foot or horseback only, and is virgin
territory for the lepidopterist. For many years following its establish-
ment in 1925 the Yosemite Field School (short title for Yosemite
School of Field Natural History) conducted annual summer excursions
into this hinterland under the capable leadership of former Park
Naturalist C. A. "Bert" Harwell. Begun as a six-day circuit of the
established Hi-Sierra Camps, this "back-country trip” developed into
an independent pack caravan that lost itself among the glaciers for a
ten-day to two-week period and seldom returned without a contribution
to the natural history of the park. During the 1933 to 1937 period the
Field School included among its members a number of students of
insects whose training and experience made it possible to observe and
record much valuable information concerning the habits of rarely en-
countered species, particularly in connection with the annual surveys of
Boundary Hill Research Reserve, instigated in 1933 by Joseph S. Dixon,
field biologist, National Park Service. Sections on butterflies in the
Research Reserve Reports were written in 1933 by John S. Garth and
Fred C. Ziesenhenne, in 1934 by Janet Mabry and Edmund D. God-
win, in 1935 by Theodore H. Eaton, Jr., and in 1937 by Robert
Godwin, Frances Cramer, and Verlin Baysinger. The 1936 report, by
R. Paul Allen and Carsten Aherns, noted succinctly: "Rain for six
days, no butterflies.” These reports are on file at the Yosemite Museum,
where they constitute an important supplement to the published report
of Garth ( 1935a) .
It was as a resurvey after an absence of 23 years from Yosemite that
the senior author undertook the present assignment at the request of
Chief Park Naturalist Douglass H. Hubbard, the intervening years
a(i):i~96, 1963
YOSEMITE BUTTERFLIES
7
having been enriched with similar experiences at Grand Canyon,
Glacier, and Grand Teton national parks. Desirous of extending the
earlier coverage of the park, he began his observations in 1956 in the
vicinity of Mather, including the lower Tuolumne, Hetch-Hetchy, and
Aspen valleys. In the same year the junior author, whose Yosemite
experience also dates from the late 1920’s, began under park auspices
an ecological study of the area adjacent to Tioga Pass, including a
transect from Pilot Peak to Mt. Dana (Tilden, 1959). Since both
investigators were limited to brief visits to the area at critical periods,
they were fortunate to find in Oakley Shields a summer resident whose
activities spanned the entire season at Tuolumne Meadows, where his
father, Allan Shields, was a seasonal naturalist. By combining records
over the succeeding 7-year period it has been possible to present a fairly
consistent picture of the situation in the park’s high country, its western
approaches, and of the Mono Basin to the east, although the coverage
of some areas is less complete than might have been desired.
THE SURROUNDING NATIONAL FORESTS
The visitor driving to and from Yosemite Valley commonly arrives
by one of three routes from the west and leaves by another. These are
California State Highways 41 via Coarsegold and Wawona, 140 via
Mariposa and El Portal (the All-Year Highway), and 120 via Grove-
land and Buck Meadows. For many miles he travels through wooded
foothills of the Sierra and Stanislaus National Forests, where oppor-
tunities for observing butterflies are equal to those within the National
Park, while collecting is unrestricted. If visiting Tuolumne Meadows,
he may wish to leave the Park by its eastern entrance, traveling
through Inyo National Forest from Tioga Pass to Lee Vining, there
joining U. S. Highway 395. Should he seek an area comparable to the
Park s high country he will find it at Saddlebag Lake, accessible by
road from Tioga Lake, and adjacent to the Hoover Wildlife Preserve
and the Hall Primitive Area. Other east-slope mountain lakes accessible
from Highway 395 are Lundy Lake, the Virginia Lakes, and the June
Lake circuit.
During the years 1950-1958 the junior writer spent considerable
time in the Mono Lake region, obtaining many records later verified
and extended by Oakley Shields. In 1958 the senior writer established
temporary residence at June Lake and from this base made one-day
trips to Gem Lake, Saddlebag Lake, and the Virginia Lakes basin. In
1957 and again in 1959 Oakley Shields crossed the Sierran Divide at
Donohue Pass and proceeded southeastward via Agnew Pass to Agnew
Meadows. In the spring of 1961 particular effort was made by Park
Naturalist Keith Trexler to explore the western foothills, where the
season begins from six weeks to two months earlier than in Yosemite
Valley. The results of these extra-limital investigations were to add a
8
GARTH AND TILDEN
/. Res. Ltpid.
number of species that rarely, if ever, occur within Park boundaries,
but which deserve consideration in any account of the butterflies of
the Yosemite region, which includes the encircling National Forest
lands as well. It is the authors’ hope that the inclusion of these locali-
ties outside the Park will serve to entice the butterfly enthusiast away
from the few crowded centers and into the less frequented but no less
rewarding byways.
YOSEMITE FOOT-TRAILS
While some of the most spectacular butterflies, including the Leto
Fritillary (Argynnis cybele leto) and the California Sister {Limenitis
bredowii calif ornica) , fly within the confines of Yosemite Valley, the
lepidopterist seeking the rarities for which the Park is famous must
be willing to do some strenuous hiking. The valley walls rise per-
pendicularly three thousand feet and more, but once their summit is
gained a large expanse of comparatively level country is accessible by
means of foot-trails that wind through alpine meadows fragrant with
wildf lowers. Over 700 miles of such trails traverse the territory within
Park boundaries, making the Yosemite a veritable entomologist’s para-
dise.
Leaving the north side of the valley floor by the Yosemite Falls
trail, a two-hour climb, best accomplished in the early morning, places
one in a position to select either the Eagle Peak, Yosemite Creek, or
North Dome trails for a day among the Parnassians ( Parnassius
clodius baldur and P. phoebis behrii) . Likewise, the ascent to Glacier
Point by the Ledge or Four-Mile trails allows the entomologist a choice
of either the Pohono or Glacier Point trails that parallel the south rim
of the valley and along which the Nivalis Copper {Lycaena nivalis)
is certain to be encountered. The "high country,’’ reached either via
Lake Tenaya or Lake Merced, is the habitat of the Ivallda Arctic
{Oeneis chryxus ivallda), Malcolm’s Checker-Spot {Melitaea damoetas
malcolmi) , and the Lustrous Copper {Lycaena cupreus) . It was the
privilege of the senior writer, as a member of the Yosemite Field
School class of 1933, to cover over 200 miles of trail outside of Yose-
mite Valley, climbing Mt. Lyell, el. 13,090, Mt. Kuna, el 12,956, and
Mt. Dana, el. 13,050 feet, on successive days, and in 1958, exactly 25
years later, to retrace a portion of this itinerary, from Tuolumne
Meadows to Merced Lake via Vogelsang Pass, with Oakley Shields.
(And aside from the trail’s being steeper and the butterflies more
elusive, things were much as he remembered them! ) Generally
speaking, June is the month for exploring the Yosemite Valley, July
for the valley’s rim, and August for the glacier country.
a(i):i-96, 1963
YOSEMITE BUTTERFLIES
9
LIFE ZONES
The incline from El Portal on the west to the Sierra Crest at the
eastern park boundary may be subdivided into five regions or life
zones, each supporting a distinctive flora and fauna. These zones, in
ascending order, are Upper Sonoran, Transition, Canadian, Hudsonian,
and Arctic-Alpine. A sixth zone, the Lower Sonoran, occurs a few miles
west in the San Joaquin Valley. Thus within the greater Yosemite
region is to be found every life zone recognized in temperate North
America with the exception of the Subtropical, found only in Florida
and southern Texas. In many cases the zones merge imperceptibly, as
when a forest predominantly of Jeffrey Pine and Incense Cedar, typi-
cally Transition, gives way to one of Red Fir and Quaking Aspen,
typically Canadian. In others the line of demarcation may be startlingly
abrupt, as when the south slope of a chaparral-clothed ridge (Upper
Sonoran) gives way to an open evergreen glade (Transition) on the
opposite-facing north slope at the same elevation. While the subject
of life zones has been exhaustively treated elsewhere, it may be said
that of the factors that determine life zones temperature is the single
most important, and that temperature is regulated by latitude, altitude,
direction of prevailing winds, proximity to bodies of water, ascending
currents of warm air from desert regions, descending drafts of cold
air from mountain tops or glaciers, character of substrate, and pattern
of drainage.
BUTTERFLIES AS LIFE-ZONE INDICATORS
Plants and animals may be divided into two groups with respect
to life zones: those that range over a wide territory and those that
adhere closely to a single zone. The former are called cosmopolites
because of their unrestricted movement; the latter are called indicators,
because their presence is considered sufficient to establish the presence
of the zone. (See GrinneU and Hall, 1919). Thus the Cony {Ochotona)
of the rock slides is an indicator of Hudsonian Zone; the Sierra Nevada
Rosy Finch {Leucosticte) of the snow banks is an indicator of Arctic-
Alpine Zone; and the Chamise {Adenostoma) of the chaparral is an
indicator of Upper Sonoran Zone. On the assumption that the most
permanent indicators are those most to be relied upon, plants and
animals might be arranged, in descending order of reliability, in the
following order: trees and shrubs, herbs and forbs, mammals, reptiles
and amphibians, and birds. In attempting to place insects in such a
scale consideration need be given to their dependence on a particular
host plant in the larval stage, their normal flight range as adults, and
their migratory habits. Thus the insect combines the fixity of the plant,
the mobility of the mammal or reptile, though to a resricted degree,
and the periodic wanderings of the bird, although less perfectly devel-
oped. The insects might with some justification be ranked between
10
GARTH AND TILDEN
/. Res. Lepid.
Figure 1. Profile of the YOSEMITE REGION showing life zones and biotic provinces, together
with a list of the butterfly species most closely restricted to them.
CALIFORNIAN PROVINCE
UPPER SONORAN ZONE
Coenonympha t. californica
Cercyonis silvestris
Argynnis c. inornata
Euphydryas c. chalcedona
Euphydryas e. rubicunda
Melitaea leanira
Apodemia m. tuolumnensis
Habrodais grunus
Strymon auretorum
Strymon a d e n o s t o m a t i s
Callophrys d. windi
Callophrys dumetorum
Callophrys iroides
Lycaena a. arota
Lycaena gorgon
Philotes speciosa
Pieris napi venosa
Epargyreus clarus
Thorybes pylades
Heliopetes ericetorum
Erynnis z. funeralis
TRANSITION ZONE
Argynnis c. leto
Argynnis z. zerene
Argynnis hydaspe
Callophrys johnsoni
Callophrys nelsoni
Everes amyntula
Colias o. chrysomelas
Thorybes diversus
Hesperia h. yosemite
Amblyscirtes vialis
CANADIAN ZONE
Argynnis irene
Boloria epithore
Euphydryas c. sierra
Melitaea hoffmanni
Lycaeides a. anna
Plebejus saepiolus
Plebejus lupini
Anthocharis s. Stella
Pyrgus ruralis
Erynnis p. lilius
Hesperia harpalus, blend
HUDSONIAN ZONE
Callophrys lemberti
Lycaena mariposa
Lycaena nivalis
Philotes b. battoides
Plebejus g. podarce
Glaucopsyche 1. Columbia
Colias behrii
Thorybes m. nevada
Polites s. tecumseh
ARCTIC- ALPINE ZONE
Oeneis c. ivallda
Euphydryas e. nubigena
Melitaea d. malcolmi
Lycaena cupreus
Lycaena p. hyp'ophlaeas
Plebejus s. comstocki
Pieris o. calyce
Parnassius p. behrii
Hesperia miriamae
ARTEMESIAN PROVINCE
CANADIAN - HUDSONIAN
Cercyonis oeta
Argynnis z. malcolmi
Lycaena heteronea
Lycaena rubidus
Lycaena editha
Hemiargus isolus
Thorybes m. nevada
Hesperia nevada
TRANSITION- UPPER SONORAN
Coenonympha t. mono
Cercyonis p. ariane
Danaus g. strigosus
Argynnis n. apacheana
Argynnis c. nevadensis
Euphydryas e. monoensis
Melitaea acastus
Limenitis w. nevadae
Apodemia m. mormo
Strymon dryope
Lycaena a. virginiensis
Everes comyntas
Leptotes marina
Philotes b. glaucon
Pieris beckeri
Colias p. hagenii
Hesperia harpalus
CALIFORNIAN \ ARTEMESIAN
YOSEMITE BUTTERFLIES
11
the plants and the vertebrates, possessing greater reliability as indica-
tors than the mammals, birds, and reptiles on which much of the eco-
logical work of the past has been done, their importance in this respect
having but lately been recognized.
The first attempts at zonal analysis of California butterflies were
of necessity incomplete and therefore inconclusive. Based upon the
record, in some cases, of but a single specimen, and without knowledge,
in many instances, of the larval food plant, the assignment of the
species to a given life zone was made without knowledge of its status
within that zone, whether as resident or vagrant. The pioneer work of
Comstock and Dammers in the 1930’s made known the life histories of
most of the butterfly species of the southern half of the state, while
workers of a younger generation, among whom may be mentioned
Don MacNeill, John Burns, G. and R. Bohart, Noel McFarland, Paul
Opler, Tom and John Emmel, and Oakley Shields, have defined the
range of species and subspecies within the entire state with exactness.
Thus the life zone normally inhabited by a butterfly of the Yosemite
region can be stated with more assurance than was possible a quarter-
century ago. A brief characterization of life zones follows:
Upper Sonoran Zone: The chaparral or "elfin forest” that clothes
the foothills from 1,500 to 4,000 feet (occasionally lower) constitutes
the Upper Sonoran Zone. It is characterized by a great variety of
shrubby plants, many of which exhibit remarkable adaptations for
conserving moisture, including anastomosing root systems that check
soil erosion. Digger Pine, California Buckeye, Poison Oak, and Chamise
{Adenostoma) are typical species, as is Interior Live Oak of the in-
cluded Foothill Woodland community. The Chalcedon Checker-Spot
{Euphydryas c. chalcedona), California Ringlet {Coenonympha tullia
calif ornica) ^ and Sylvan Satyr {Cercyonis silvestris) seldom stray be-
yond the confines of this life zone.
Transition Zone: Between the Austral zones (Lower and Upper
Sonoran) and the Boreal zones (Canadian, Hudsonian, and Arctic-
Alpine) lies a broad intermediate area known as Transition Zone. Its
coniferous forest contains the trees of greatest commercial value: Big
Tree, Yellow Pine, Jeffrey Pine, Sugar Pine, Incense Cedar, White
Fir, and Douglas Fir. Azalea, Nuttall Dogwood, and Black Cotton-
wood fringe its streams; Black Oak and Golden Cup Oak clothe its
valley floors and mountainsides, respectively. The Leto, Hydaspe, and
Zerene fritillaries {Argynnis cybele leto, A. hydaspe, and A. z. zerene)
represent its considerable butterfly population. Transition Zone ex-
tends from 4,000 (or less in some localities) to 7,000 feet, merging
imperceptibly into Canadian Zone along its upper border.
Canadian Zone: Red Fir replaces White Fir in the open forest and
Quaking Aspen displaces Azalea and Black Cottonwood along the
water courses. A secondary chaparral, reminiscent of Upper Sonoran
Zone, but composed of matted Snow Brush {Ceanothm) ^ Chinquapin
12
GARTH AND TILDEN
/. Res. Lepid.
{Castanopsis) , and Huckleberry Oak {Quercus vaccinifolia) , covers
the steeper slopes. The Western Banded Elfin {Gallop hrys eryphon) ,
Sierra Checker-Spot {Euphydryas chalcedona sierra)., and Western
Meadow Fritillary (Boloria epithore) are good butterfly indicators.
Canadian Zone extends from 7,000 to 9,000 feet (or less in some
localities), with Lodgepole Pine invading from its upper margin.
Hudsonian Zone: At about the 8,500 foot level Lodgepole Pine
appears, to be joined at the 9,000 foot level by Mountain Hemlock to
form the Hudsonian forest, discontinuous because of the tremendous
rock slides and glacial cirques. Tuolumne Meadows, at 8,600 feet, is
pure Hudsonian and here are found Behr’s Sulphur {Colias hehrii) ,
the Podarce Blue (Plebejus glandon podarce), and the Mariposa Cop-
per {Lycaena mariposa) . The upper limits of Hudsonian Zone are
defined by timberline at approximately 11,000 feet.
Arctic- Alpine Zone: The Alpine Willow, stunted to a height of a
few inches, and the Sierra Nevada Rosy Finch inhabit the bleak talus
slides and snow banks above timber line. The zone is characterized by
perennial herbs with small tops and large root systems. The heathers,
Bryanthus and Cassiope, and Alpine Sorrel (Rumex) cling to meager
patches of damp soil. The Ivallda Arctic {Oeneh chryxus ivallda),
Malcolm’s Checker-Spot {Melitaea damoetas malcolmi) , and the tiny
Yosemite Blue {Plebejus shasta comstocki) fly from 11,000 feet (occa-
sionally less) to the tops of Mt. Dana and Mt. Lyell. Recently Miriam’s
Skipper {Hesperia miriamae) has been added to the strictly Arctic-
Alpine list.
It should be remembered that it was in California, and particularly
on the west slope of the Sierra Nevada, that most of the detailed work
on life zones, including the delineation of indicator species, was done.
It is also on the west slope of the Sierra Nevada and Cascade ranges
that life zones attain their optimum development, occurring in regular
succession from west to east in response to moderate gradients of
slope and a comparatively mild, ocean-controlled climate. Elsewhere
life zones are likely to occur haphazardly, with extreme local conditions
often overriding the effects of latitude and altitude, which elsewhere
are the principal factors governing temperature, on which the life-
zone concept was originally based. One of these, the humidity factor,
was taken into consideration by Grinnell, Storer, and Dixon in both
the Yosemite and Lassen sectors, which included an arid region east
of the Sierra divide. Their concept of a trans-montane fauna con-
ditioned by high aridity as opposed to a cis-montane fauna con-
ditioned by high humidity leads us to the principle of replacement of
one set of zonal indicators by another whenever certain natural boun-
daries are transgressed. This is better expressed within the context of
a more recent ecologic concept, that of the Biotic Province.
2(i):t-^6, 1963
YOSEMITE BUTTERFLIES
13
BIOTIC PROVINCES
The replacement of one set of indicator species by another can be
so complete that we may question whether we are dealing with the
same life zone in two widely separated regions, or even in adjacent
regions, when each indicator species has been replaced by a different
but closely related species or subspecies. The solution to this predica-
ment is found in the concept of the Biotic Province, originated by
Vestal (1914) and developed by Dice (1943), who divided the North
American continent into 28 biotic provinces, of which 14 occur in
states west of the Rockies, 7 in California. These are geographical
areas, permanent and mutually exclusive, and containing within their
confines vertical bands that Dice calls, not life zones^ but life belts. In
such a system the principle of replacement need not be considered, for
by definition no life belt extends beyond its province, although similar
life belts may occur in adjacent provinces.
In the Yosemite region, as has been mentioned, all life zones are
represented from Lower Sonoran (in the San Joaquin Valley) to
Arctic-Alpine, with a correspondingly wide spectrum of plant and
animal life. Park boundaries are drawn, however, in such a manner that
the eastern boundary coincides with the crest of the Sierra Nevada,
thus limiting the park proper to one biotic province, the Californian.
If we extend the Yosemite sector eastward to include the eastern slope
of the Sierra (as did Grinnell and Storer in extending their Yosemite
sector to the shore of Mono Lake), we will have added another biotic
province, the Artemesian, and with it the replacement species of that
province, potentially doubling the list of species for zones above
6,500 feet. No more than this could be accomplished by combining the
species lists from two different parks, the one completely Californian,
the other Artemesian. Furthermore, such an extension is necessary in
order to compare the butterfly faunas of Yosemite and Lassen, whose
boundaries stop at the Sierran Crest, with those of Glacier and Rocky
Mountain, whose boundaries include territory on both sides of the
Continental Divide.
An unexpected dividend for the biologist of this meeting of faunas
along the border of two biotic provinces is the opportunity of studying
natural hybridization. Perhaps no better experimental material has
been provided by nature for this purpose than the butterflies known
as Admirals (genus Limenitis') . These are found throughout North
America and are known as allopatric, or replacement, species. Oppor-
tunity to observe natural hybridization between Californian L. lorquini
and Artemesian L. weidemeyerii nevadae occurs where their ranges
meet on the west shore of Mono Lake. A similar opportunity to observe
natural hybridization between Montanian L. lorquini hurriaoni and
Saskatchewan L. arthemis rubrofasciata occurs on the east side of
Glacier Park, at Many Glacier. Whether the fact that they hybridize
readily reduces the Limenitis complex to a single polymorphic species
14
GARTH AND TILDEN
/. Res. Lepid.
is a debatable question; however, it should be noted that the interbreed-
ing is limited to a narrow buffer zone and does not extend to entire
populations, and that wherever it occurs both parent species are pres-
ent, as would be required to perpetuate the hybrid, which is presumed
to be sterile.
PLANT COMMUNITIES
A more subtle ecologic division than either life zone or biotic prov-
ince has long been sought by entomologists attempting to define more
closely the habitats of insect species. This has at last been provided
in the system of plant communities developed by Munz and Keck
(1949, 1950 ). Of the 28 California plant communities recognized by
these authors, at least 16 (plus two of our own designation) occur in
the greater Yosemite region. Here an attempt is made to list in se-
quence the plant communities that one might encounter in crossing the
Sierra Nevada from San Joaquin Valley to Mono Lake, with a side trip
into Yosemite Valley:
1. Valley Grasslands (on leaving the Valley) .
2. Foothill Woodland ( as around Cathay) .
3. Chaparral (the familiar "elfin forest”).
4. Coastal Sage Scrub (formerly California Sagebrush Associa-
tion, includes Honey Sage Scrub, occurs sporadically).
5. Oak Woodland (including Golden Cup Oak lands; as Yose-
mite Valley).
6. Mixed Evergreen Forest (Laurel-Madrono association; as
Yosemite Valley walls).
7. Yellow Pine Forest (monotypic, with an understory of
"Mountain Misery”).
8. Mixed Forest (non-pure Douglas Fir, Yellow Pine, and broad-
leaf trees below pure conifers). Our designation; probably
an ecotone.
9. White Fir-Cedar Forest (on cut-over land, as at Mather).
Our designation; a successional sub-climax or post-climax
community.
10. Red Fir Forest (as between Tamarack Flat and Smoky Jack).
11. Lodgepole Pine Forest (as Tuolumne Meadows).
12. Subalpine Forest (often with Mountain Hemlock and White-
Bark Pine; as Tioga Pass) .
13. Alpine Fell-Fields ("rock gardens” above timber line; as
Gay lor Lakes).
14. Juniper Woodland (eastern declivity; as below Tioga Pass).
15. Pihon- Juniper Woodland (lower down; as around Lee Vin-
ing).
16. Sagebrush Scrub (Great Basin Sage, Artemesia tridentata) .
17. Shadscale Scrub (the Salt-Bush type of scrub).
18. Alkali Sink (as around Mono Lake) .
2(t):i-^6, 1963
YOSEMITE BUTTERFLIES
15
Figure 2. Map showing the YOSEMITE high country, indicating roads . trails. ..........
streams , areas over 11, 000 feet in elevation and collecting localities:
1933-1934 : 3, Mt. Dana; 6, Florence Lake; 8, Glen Aulin; 9, Kuna Crest; 12, Mt Lyell; 18, Cold-
water Canyon.
1956-1962; 59, Agnew Pass; 60, Bert L.; 61, Cockscomb Pk. ; 62, Mt. Dana, N. slope; 63, Dog L. ;
64, Donohue Pass; 65, Elizabeth L.; 66, Gaylor L. ; 67, Glen Aulin; 68, Helen L. ;
70, Lembert Dome; 71, Lyell Base Camp; 72, Lyell Fork Meadow; 74, Mammoth Pk. ;
75, Merced R.; 76, Mono Pass; 77, Pilot Pk. ; 78, Rafferty Ck. Tr.; 79, Saddlebag
80, Tioga Pass; 81, NW above Tioga Pass; 83, Thousand Island L.; 84, Tuolumne
Meadow; 85, Unicorn Pk.; 86, Vogelsang Pass.
East slope , 1956-1962: 92, Gull L.; 93, June L.; 94, LeeVining Cr. ; 95, Mono L. ; 98, Warren Ck.
16
GARTH AND TILDEN
/, Res. Lepsd.
[One of US (Tilden) suggests the presence of still another
east-slope brush community, consisting of Cercocarpus, Pur-
shia, and desert Pr7mus, not provided for in the system pro-
posed by Munz and Keck.]
It will be noted that by far the greater number of plant communi-
ties ( 1 3 as against 5 ) are found on the western slope, which has the
greatest extent both in distance and in elevation. It will further be
seen that as one progresses beyond 13, which represents the Sierran
crest, the plant communities are entirely different from those before
13, yet they occupy corresponding altitudinal belts and life zones in
reverse order. This difference in plant life is paralleled by a difference
in animal life, including the insects, and is evidence that each slope lies
in a different biotic province, where relative humidity, rather than
temperature difference based on altitude, is the controlling factor. Per-
haps no better example could be given of the need for both systems
(life zones and biotic provinces) .
FOOD PLANTS
The lepidopterist who pursues his interest seriously becomes in-
creasingly aware of the dependence of insects on plants. For not only
do butterflies as adults seek flowers for nectar, but as caterpillars they
depend on vegetation for sustenance. Many are highly selective in their
choice of plant food, adhering to one kind and rejecting all others.
More frequently, however, any one of a group of related plants will
serve: witness the predilection of the Whites for mustards, or of the
Sulphurs, for legumes. The lepidopterist soon learns to equate these
butterfly preferences with plant groups possessing certain structural
resemblances, particularly as regards their flowers, and recognized by
botanists as plant families. A list of thirty-eight common plant families
and one hundred eleven of their genera on which butterflies of
the Yosemite region are known or believed to feed as larvae is pre-
sented with the hope that it will lead to further investigation.
Occasionally, and for no apparent reason, a butterfly will be found
to feed upon a plant quite unrelated to those usually preferred by its
group. Thus Colias hehrii is known to be a Vaccinium feeder, whereas
most .species prefer legumes. But as Hovanitz (1950) has
shown, the V accinmm-iGQ6.mg habit is shared by several Colias species
in other parts of the world. Since it is highly improbable that such a
choice of food plant would have been made independently by each of
them, this food habit becomes a clear indication of relationship, setting
the Vaccinium feeders apart from others of their genus. When one of the
Whites, Neophasia menapia, forsakes the mustards usual to its group
for the needles of a pine tree, we may be certain that it is not tempor-
arily off its diet, but that it is following a behavior pattern as old as the
genus itself, and perhaps responsible for its segregation.
2(i):i-9S, 196}
YOSEMITE BUTTERFLIES
17
Often a butterfly named for the plant believed to be its host has
proven to feed upon another. The hair-streak, Strymon adenostomatis,
was found to feed on Mountain Mahogany (Cercocarpus) , and not on
Chamise {Adenostoma) , as its given name would imply. (It should
have been called cercocarpivofous”\ ) Indeed, the question as to wheth-
er the Blues, Plebejus acmon 3.nd P. lupini, deserve specific distinction
may ultimately rest upon whether they feed exclusively on Lupine
(Lupinus) and on Buckwheat {Eriogonum) , respectively, or indis-
criminately on both.
Yosemite butterflies for which the larval food plant is either un-
known or in need of confirmation are the following:
Unknown
Melitaea damoetas malcolmi
Melitaea hoffmanni
Callophrys doudoroffi windi
C allop hr ys lemberti
Lycaena nivalis
Plebejus glandon podarce
Plebejus shasta comstocki
Lycaeides ar gyro gnomon anna
Thorybes mexicana nevada
Thorybes diver sus
Erynnis persius
Erynnis propertms
Polites sonora
Needing Confirmation
Callophrys nelsoni (Libocedrus?)
Lycaena rubidus ( Rumex? )
Lycaena phlaeas hypophlaeas
( Rumex? )
Plebejus lupini (Eriogonum?)
Philotes speciosa {Eriogonum
in Yosemite.^)
Colias zerene {Amorpha not
found in Yosemite)
Parnassius clodius baldur
(for Vaccinium only)
Pyrgus ruralis ( for Potentilla
— Horkelia only)
Erynnis pacuvius lilius
( Ceanothus? )
A documented observation of oviposition (egg laying) of any of
these species, with positive identification of the plant of choice, or
better still, the rearing of any of them from egg to adult would be an
important contribution to the natural history of the Yosemite region.
SOME PLANTS ON WHICH YOSEMITE BUTTERFLIES
FEED AS LARVAE
In the left hand column of the list ( Table 1 ) that follows, host
plants are listed alphabetically by family and genus. (To have listed
plants by species would have prolonged the list unduly.) In the
middle column the common name of the plant is given. In the right
hand column butterfly species known or believed to feed on that
plant are listed alphabetically by genus and species. (Subgenera are
omitted, as are subspecies, where more than one occurs in Yosemite,
unless each is known to have a different food plant.) The symbols o, 1,
18
GARTH AND TILDEN
/. Res. Lepid.
and r indicate that the butterfly has been seen to ovipost on, the larvae
have been taken from, or the insect reared to maturity on that plant
by one of us. Otherwise, names of host plants have been taken from
the literature, carefully culled to eliminate genera not known to occur
in the Sierra Nevada. Butterfly species feeding on more than one plant
are starred*.
Grasses (POACEAE=GRAMINACEAE) and Sedges (CYPER-
ACEAE) are not listed because detailed information concerning the
genera involved is lacking. Known to be grass feeders are all the
SATYRIDAE and, of the HESPERIIDAE, the subfamily Hesperiinae.
It is believed that when this information becomes available, it will be
shown that butterflies exhibit as marked a preference for certain of
the monocotyledonous grasses and sedges as they do for certain of the
dicotyledonous plants.
THE GEOLOGICAL BACKGROUND
The landscape of the Yosemite region has been moulded by many
forces, but by far the most impressive of these has been glaciation.
The work of glaciers may be observed in the U-shaped gorges of the
Tuolumne and Merced rivers, in the hanging valleys of Little Yosemite
and Bridal Veil Creek, in the polished domes and erratic boulders, in
the glacial lakes (called tarns) and cirques, and in the minaret summits
of Cathedral and Unicorn peaks. Less evident, but no less genuine, are
the effects of glaciation on the flora and fauna, including the butter-
flies. For as the glaciers retreated, the cold-adapted species followed
them northward and to the mountain tops, where small glaciers con-
tinue to exist on the shaded slopes of Lyell, Maclure, Dana, and Con-
ness. Thus the entire upper tier of life zones, the Canadian, Hudsonian,
and Arctic-Alpine, is inhabited by butterfly species whose nearest rela-
tives fly either on other isolated mountain tops in temperature regions
or at near sea level in Canada, Hudson Bay, or the Arctic regions, as
these zonal names imply. This discontinuous "boreo-alpine” distribu-
tion is as much a consequence of Pleistocene glaciation as the polished
granite over which one walks along the Merced Lake trail, or the
lateral moraines that rim the montane meadows.
A second consequence of Yosemite’s geological history is that those
butterflies that now fly above timberline, and in particular the grass-
feeding Satyridae, such as The Ivallda Arctic {Oeneis chryxus ivallda)
and Riding’s Satyr {Eumenis ridingsii) , and Hesperiinae, such as the
newly found Miriam’s Skipper {Hesperia miriamae) , having no trees
but only barren rock on which to alight, have come to resemble exactly
the color, shade, and texture of the particular rock found in their re-
stricted habitat, whether it be sedimentary or metamorphic. (See
Hovanitz, 1940). In order that the lepidopterist may better appreciate
these earthen pigments that provide the backdrop for the Yosemite
1963
YOSEMITE BUTTERFLIES
19
landscape and constitute the palette from which the colors of Yose-
mite’s alpine butterflies (Plate III) have been selected, although with-
out design on the insect’s part, the following quotation by Francois E.
Matthes, father of Sierran geology, is given. The italics are the authors’.
"Next to the broad belt on the lower slope of the Sierra Nevada, the
masses of metamorphic rocks situated near the crest of the range are the most
extensive. They make up the bulk of Mount Dana, Mount Gibbs, and Parker
Peak, as well as the jumble of mountains north of Tioga Pass, whose central
summit is Mount Warren. These mountains, in consequence of their compo-
sition, are variously tinted in subdued yellotvs, browns, reds, and purples and
by contrast with the pale-gray peaks of granite near by appear somber, as if
overcast by perpetual shadows. The metamorphic rocks in this crest region,
however, differ appreciably in character as well as in structure from those
in the lower belt. There is but little slate among them, schist, quartzite, and
volcanic rocks being predominant; and the folding is less deep and less
complex. These facts are readily observed on and about Mount Dana, which
is capped by gently flexed beds of volcanic origin.”
— Matthes, 1930, p. 26.
ACKNOWLEDGMENTS
Among agencies and individuals cooperating in the preparation
of this report should be mentioned the National Park Service, Depart-
ment of Interior, and in particular Park Superintendent John C. Pres-
ton and Chief Park Naturalist Douglass H. Hubbard, Yosemite
National Park, who issued the necessary permits, and Park Naturalist
Keith Trexler, who assisted with the collecting. Among those provid-
ing suggestions and information on technical matters were John Burns,
Harry Clench, William Hovanitz, C. Don MacNeill, Paddy McHenry,
and Paul Opler. Among those furnishing advance information concern-
ing manuscripts in preparation were Cyril F. dos Passos, C. Don
MacNeill, and Paul and Anne Ehrlich. Among museums and curators
whose collections were consulted were the California Academy of
Sciences, Edward S. Ross, and the Los Angeles County Museum, Lloyd
M. Martin. Finally, two investigators whose efforts so splendidly suj>-
plemented our own, and whose records and observations we have used
to such advantage, were Edmund D. Godwin and Allan Oakley Shields.
REGARDING NAMES USED IN THIS PAPER
Since the publication of "Butterflies of California” (Comstock,
1927), no comparable book on California butterflies has appeared.
The excellent color plates make this a most useful reference, but time
and the activities of butterfly students have brought about many
changes. Additional species have been found to occur in California;
numerous subspecies have been described; and many revisionary works
have appeared. Among these may be mentioned those on Ringlets
(Davenport, 1941; Brown, 1955), on Fritillaries (dos Passos and
20
GARTH AND TILDEN
/. Res. Lepid.
Fig, 3. Mt. Lyell, el. 13,900 ft., from tree-line at about 11,000 ft.
(Upper Lyell Base Camp). Arctic-Alpine life zone, Alpine Fell-Fields, with
montane meadow and rock garden associations. Habitat of Lycaena phlaeas
hypophlaeas, Euphydryas editha nubigena, Melitaea damoetas malcolmi, and
Oeneis chryxus ivallda. — Joseph S. Dixon.
2(i);i-96, 1963
YOSEMITE BUTTERFLIES
21
Grey, 1947), on Checker-Spots (Gunder, 1929; Bauer, 1961), on
Metal-Marks (Opler and Powell, 1961), on Hair-Streaks (Ziegler,
I960; Clench, 1961), on Coppers (Klots, 1936; Clench, 1961), on
Blues (Nabokov, 1949; Downey, 1961), and on Skippers (Lindsey,
Bell, and Williams, 1931; Lindsey, 1942). As a result, only a fraction
of our California butterflies continue' to bear the generic and specific
names by which they were known as recently as 1927.
Following the publication of "Check List of Lepidoptera of Canada
and the United States of America” (McDunnough, 1938), incorpor-
ating changes recommended in "Generic Names of Holarctic Butter-
flies” (Hemming, 1934), together with the findings of many other
workers, no similar standard of reference has been advanced. Publica-
tion of a new check list by C. F. dos Passos is believed imminent, and
changes made since 1938 will be reflected therein. Many of these will
have been anticipated in "A Field Guide to the Butterflies” (Klots,
1951 ), but this admirable little book treats only the eastern species.
Still other modifications appear in "How to Know the Butterflies”
(Ehrlich and Ehrlich, 1961 ) .
In deciding what names to use in this article, the authors have tried
to steer a moderate course, adopting many names now accepted by
common usage, but avoiding some of the more recent innovations until
these shall have met with general approval. The sequence and arrange-
ment of genera and families follows as closely as possible what we are
led to believe will be the arrangement of the forthcoming check list
already referred to. The more advanced student as well as the pro-
fessional should be able to make use of that publication in conjunc-
tion with this one, which is intended primarily as a field guide, and
not primarily as a contribution to the systematic literature.
The authors’ departure from a strict interpretation of the Rules of
Zoological Nomenclature in designating the western subspecies of
AtUdes halesus as estesi Clench rather than as corcorani Gunder is de-
liberate. For while it is conceded that Gunder’s name is the older of
the two, it was applied, not to the entire race, but to a single aberrant
individual. We believe that intent should be given due weight in such
matters, and it was certainly not Gunder’s intent to define the western
subspecies, whereas it was Clench’s. The same logic is followed in
attributing Lycaeides melissa inyoensis to Nabokov, and not to Gunder.
22
GARTH AND TILDEN
/. Res. Lepid.
ACCOUNT OF SPECIES OF THE YOSEMITE REGION
SATYRIDAE
la. Coenonympha tullia californica West. & Hew. PI. I, fig. c
b. Coenonympha tullia mono Burdick PI. IV, fig. a
THE CALIFORNIA RINGLET and THE MONO RINGLET
are the two races of this widespread species found in the Yosemite
region, the former flying in the western foothills, the latter on the
sage-brush flats of Mono Basin. Weak fliers, they keep close to the
grasses on which the larvae feed. Following the studies of Davenport
(1941) and Brown (1954, 1955), all North American ringlets except
the rare Hayden’s Ringlet, Coenonympha haydenii (Edw.), of Wy-
oming and adjacent states are considered subspecies of the Eurasian
C. tullia (Muller), although Burdick (personal communication) was
inclined to regard his mono as a derivative of the Great Basin-Rocky
Mountain C. t. ochracea Edw., which he appeared to regard as a dis-
tinct species.
Life Zones: Upper Sonoran (Transition). Plant Communities: Val-
ley Grasslands, Foothill Woodland; Sagebrush Scrub, Marsh. Host Plants:
POACEAE (Grasses).
2. Cercyonis pegala ariane (Bdv.)
It is with some hestitation that the name THE ARIANE SATYR
is applied to specimens from Mono Lake, because it is BARON’S
SATYR, Ceifcyonis pegala haroni (Edw.), that occurs in a similar
situation in northeastern California and southeastern Oregon.
Life Zones: Upper Sonoran. Plant Communities: Pihon-Juniper
Woodland, Marsh, and Montane Meadow. Host Plants: POACEAE
(Grasses).
3. Cercyonis silvestris (Edw.) PI. I, fig. e
THE SYLVAN SATYR is a creature of the chaparral forest that
clothes the foothills. A lover of shade, it seeks to lure the pursuer ever
deeper into the undergrowth. Its favorite haunts are groves of Man-
zanita. Scrub Oak, and Mountain Mahogany, interspersed with an occas-
ional Digger Pine.
Life Zones: Upper Sonoran (Transition). Plant Communities:
Foothill Woodland, Chaparral. Host Plants: POACEAE (Grasses).
4. Cercyonis oeta (Bdv.) PI. IV, fig. c
THE LEAST SATYR was not encountered in Yosemite Park
during the 1933-1934 period, although it was known to inhabit the
i$6j
YOSEMITE BUTTERFLIES
23
dry eastern slope of the Sierra Nevada. Since that time, however, it
has been found on several occasions in the Tioga Pass region, but
always within sight of the Sierran Divide.
Life Zones: Transition, Canadian, Hudsonian ( Arctic- Alpine ) .
Plant Communities: Juniper Woodland, Pihon-Juniper Woodland,
Sagebrush Scrub (Alpine Fell-Fields). Host Plants: POACEAE
(Grasses) .
5. Oeneis chryxus ivallda (Mead) PI. Ill, fig. b
THE IVALLDA ARCTIC inhabits the bleak, wind-swept sum-
mits that culminate in Mt. Lyell, Mt. Dana, and Mt. Conness. They
may be encountered in a boulder-strewn clearing between stands of
Mountain Hemlock, as near Vogelsang Camp, on barren ridges, as the
crest above Tioga Pass crossed by the Gaylor Lakes trail, or on the
moraines just east of Tioga Pass. Edmund Godwin reported "many
seen at 12,000 feet on Matterhorn” in 1934. Erratic fliers, extremely
difficult to capture, the Arctics have a habit of leaning with the wind
when alighting, their mottled wings resembling a patch of lichen.
According to Hovanitz (1940), the gray forms are found on granite,
the brown forms on sedimentary rock. The line of contact between
the two formations passes through the Yosemite sector and is a con-
spicuous feature of Sierran geology. Because of the inaccessibility of
its habitat, the IVALLDA ARCTIC presents the ultimate challange to
the energetic lepidopterist.
Life Zones: Arctic- Alpine (Hudsonian). Plant Communities: Al-
pine Fell-Fields, Subalpine Forest (Glacial Moraine). Host Plants:
POACEAE ( Grasses ) .
DANAIDAE
6. Danaus (Danam) plexippus (Linn.)
THE MONARCH is common in Yosemite Valley and throughout
the lower elevations of the park. Various species of Milkweed grow
along the Merced River at Old Village, at Mirror Lake, and at the
fork in the road to the Giant Yellow Pine, as well as along the higher
trails that follow the canyon rim. It also occurs on the eastern Sierra
slope, at Mono Lake, and has been found at Upper Lyell Base Camp
( Shields ) . The banded larvae are easily discernible, but the waxy green
chrysalids defy detection.
Life Zones: Unrestricted below Hudsonian. Host Plants: ASCLE-
PIADACEAE: Asclepias (True Milkweeds).
7. Danaus (Tasitia) gilippus strigosus (Bates)
THE STRIATED QUEEN in another milkweed feeder that occurs
in Yosemite territory only on the dry east side of the Sierra Nevada,
in the vicinity of Mono Lake. It may be distinguished from Danaus
{Tasitia) gilippus berenice (Cram.) of the southeastern United States
by the scattered white scales along the wing veins. Both are sub-
species of the tropical D. (T. ) gilippus (Cram.), ranging northward
from Brazil.
24
GARTH AND TILDEN
/. Res. Lepid.
Life Zones: Sonoran, both Lower and Upper (= Austral). Plant
Communities: Many. Host Plants: ASCLEPIADACEAE: Asclepias
(True Milkweed), and several other milkweed genera.
NYMPHALIDAE
S'. Argy finis {Semnopsyche) cybele leto ISeht:
THE LETO ERITILLARY is the handsomest butterfly of the
Yosemite Valley and the species most certain to attract the attention
of the park visitor in the late summer season. Erom the thistles at
Mirror Lake, which they visit for nectar, the butterflies may be traced
to the sequestered meadow near Camp 9. Here the velvet-brown females
deposit their eggs among the dried grasses, while the ruddy males hover
above them, for leto is one of the two California fritillaries in which
the sexes differ in color, the other being the following apacheana.
Life Zones: Transition. Plant Communities: Poplar Woodland
(Black Cottonwood). Host Plants: VIOLACEAE: Viola (Violet).
9. Argynnis {Speyeria) nokomis apacheana Skin.
PL IV, fig. k
THE APACHE ERITILLARY, athough of another species, may
be considered the eastern Sierran counterpart of The Leto Eritillary of
the western Sierra. Even more spectacular than leto, it flies in the well
watered meadows of the Owens River drainage, and has been found
recently at Gull Lake and Mono Lake. The females emerge a full two
weeks later than the males, and so dissimilar are the sexes in color
that they might be mistaken for different species. The under side of
the male is illustrated in color.
Life Zones: Upper Sonoran (Transition). Plant Communities: Wet
Meadow, Marsh. Host Plants: VIOLACEAE: Viola (Violet).
10a. Argynnis {Speyeria) zerene zerene Bdv. PL I, fig. b
b. Argynnis {Speyeria) zerene malcolmi Comst.
THE ZERENE ERITILLARY is common along Highway 41 from
Chinquapin to Wawona, in the Mariposa and Tuolumne Big Tree
groves, and at Mather and Hetch-Hetchy summit. Every patch of
thistle will bear investigation, for this fritillary is highly susceptible
to its lure. Its Sierran east-slope counterpart, MALCOLM’S FRIT-
ILLARY, is found along the road from Highway 395 to Mammoth,
and at Warren Creek on the Tioga Road. Its first choice for nectar is
Monardella, a dwarf mint.
Life Zones: Transition, Canadian. Plant Communities: Yellow
Pine, White Fir (Red Fir), Mixed Coniferous forests for A. z. zerene,
Sagebrush Scrub, Jeffrey Pine Forest, Pihon-Juniper, and Juniper
Woodland for A. z. malcolmi. Host Plants: VIOLACEAE: Viola
(Violet).
11a. Argynnis {Speyeria) callippe inornata Edw.
b. Argynnis {Speyeria) callippe nevadensis Edw. PL IV, fig. b
THE PLAIN ERITILLARY flies in the western foothills of the
Yosemite region in the late spring. It has been found at Fish Camp,
2(t):i‘p6, 196}
YOSEMITE BUTTERFLIES
25
Fig. 4. Research Reserve, el. 8,700 ft. Open slope with Western White
Pine predominant; Lodgepole Pine and Red Fir also present, with understory
of Manzanita and Huckleberry Oak. Canadian life zone. Philotes battoides
and P. shasta comstockii hover over the dwarf buckwheat and sparse grasses
ruralis may be seen here.—Neva Snell.
26
GARTH AND TILDEN
/. Res. Lepid.
at Briceburg, and at Indian Flat, below El Portal. THE NEVADA
FRITILLARY, its eastern Sierran counterpart, flies in the Mono Lake
region in early summer. It has been found at the summit above Crowley
Lake on Balsam Root ( Balsamorhiza ) , a large-leaved composite. The
Great Basin races of callippe tend to be green and silvered below, the
races of the western foothills brown and unsilvered.
Life Zones: Upper Sonoran (lower Transition). Plant Commun-
ities: (Chaparral) Foothill Woodland, Yellow Pine and Mixed Forest
for A. c. inornata; Sagebrush, Pihon-Juniper Woodland, Jeffrey Pine
Forest openings. Juniper Woodland for A. c. nevadensis. Host Plants:
VIOLACEAE: Viola (Violet).
12. Argy finis {Speyeria) egleis Behr
Syn. Argynnis {Speyeria) montivaga Behr
THE EGLEIS FRITILLARY, known for many years as THE
MOUNTAIN VAGABOND, is a species of uniformly small size
characterized by a slight thickening of the veins of the primaries. It
flies with The Arge Fritillary at Tioga Pass and is common at all ele-
vations above 6,000 feet, preferring the minted slopes, whereas A.
mormonia arge prefers the meadows. On the west slope of Mt. Dana
it fairly swarms in the late afternoon sunlight.
Life Zones: upper Transition, Canadian, Hudsonian (Arctic-Al-
pine). Plant Communities: Openings in Mixed Coniferous, White
and Red Fir, and Lodgepole Pine Forests (Alpine Fell-FieldsJ . Host
Plants: VIOLACEAE: Viola (Violet).
13. Argynnis {Speyeria) atlantis irene Bdv. PL 11, fig. h
THE IRENE FRITILLARY of Yosemite, as determined for the
earlier survey by Dr. J. McDunnough, is a dark race, some specimens
of which are strongly suggestive of The Hydaspe Fritillary. Irene,
however, is found at elevations well above those at which hyaspe
occurs, and for this reason should not be confused with it. One of us
(Tilden) has taken it also on the Sonora Pass Road, between Straw-
berry and the Clark Fork Road. In Yosemite it is something of a rarity.
Life Zones: upper Transition, Canadian. Plant Communities: Yel-
low Pine-Sugar Pine, White Fir and Red Fir Forests. Host Plants:
VIOLACEAE: Viola (Violet).
14. Argynnis {Speyeria) hydaspe Bdv.
THE HYDASPE FRITILLARY first appears in Yosemite Valley
about the Fourth of July, flying rapidly across the valley floor but paus-
ing over the Monardella of the talus slopes. It is completely unsilvered
and the creamy spots on the under side give a "checker-board” effect
quite different from the purplish shades and frequent light silvering of
the somewhat similar but usually larger zerene.
Life Zones: Transition, lower Canadian. Plant Communities: Yel-
low Pine, Mixed Coniferous, and White Fir Forests. Host Plants:
VIOLACEAE: Viola (Violet).
I $6}
YOSEMITE BUTTERFLIES
27
15. Argynms (Speyeria) mormonia arge Stkr. PI. Ill, fig. 1
The Sierran race of THE MORMON FRITILLARY flies with THE
EGLEIS FRITILLARY throughout the park. Males lack the enlarged
veins of the forewing; females are more finely marked than those of
egleis; otherwise the two species are indistinguishable, and particularly
so when on the wing. The active males range widely, but the females
will be found near moist meadows. The Rocky Mountain race of A.
(S.) mormonia is eurynome Edw.
Life Zones: Canadian, Hudsonian ( Arctic-Alpine) . Plant Com-
munities: Sub-alpine Meadows; Alpine Fell-Fields, Meadow and Grass-
land. Host Plants: VIOLACEAE: Viloa (Violet).
16. Boloria {Clossiana) epithore (Edw.) PL II, fig. d
THE WESTERN MEADOW FRITILLARY favors small, open
glades beneath fir trees. Along Yosemite Creek it is particularly abund-
ant in mid-July. There is also a colony on Dana Trail, where it passes
through a marsh before rising sharply onto the slope of the mountain.
The under side of the secondaries resembles Boloria ( C. ) toddi ( Holl. ) ,
a widely ranging eastern species; however, in epithore the outer margin
of the primaries is rounded, whereas in toddi it appears square-cut.
Life Zones: Canadian (Hudsonian). Plant Communities: White
and Red Fir Forest; Bogs and Marshes. Host Plants: VIOLACEAE:
Viola (Violet).
17a. Euphydryas chalcedona chalcedona (Dbldy. & Hew.)
b. Euphydryas chalcedona sierra (Wgt.)
THE CHALCEDON CHECKER-SPOT ranges throughout the
foothills of the western portion of the state, being replaced in the
Sierran highlands by THE SIERRA CHECKER-SPOT, now recog-
nized, through the work of J. D. Gunder (1929), as a subspecies,
or geographical race. Typical chalcedona may be found abundantly
among the chaparral-clothed foothills, whereas race sierra prefers the
wooded glades of the evergreen belt. A variety of food plants is
acceptable to the voracious larvae.
Life Zones: Upper Sonoran for E. c. chalcedona; Canadian for E. c.
sierra. Plant Communities: Foothill Woodland, Chaparral for E. c.
chalcedona; Fir Forest for E. c. sierra. -Host Plants: SCROPHULAR-
lACEAE: Scrophularia (Bee Plant); Pentstemon ( Pentstemon ) ;
Castilleja (Paint Brush); Mimuh/s (Monkey Flower); and Diplacus
(Sticky Monkey Flower) for E. c. chalcedona; BOR AGIN ACE AE:
Mertensia ciliata var. stomatechoides (Mertensia), for E. c. sierra, ac-
cording to Davenport and Dethier (1937) CAPRIFOLIACEAE:
Symphoricarpos albus (Snow Berry) for E. c. chalcedona.
18a. Euphydryas editha ruhicunda (Hy. Edw.) PL I, fig. d
b. Euphydryas editha nuhigena (Behr) PL III, fig. c
c. Euphydryas editha monoensis Gund. PL IV, fig. f
THE RUDDY CHECKER-SPOT flies in the western Sierra from
28
GARTH AND TILDEN
/. Res. Lepid,
Tulare to Sierra counties, its "lowland” counterpart of the eastern
Sierra being THE MONO CHECKER-SPOT. As suggested by Com-
stock (1927), and confirmed by Gunder (1929), THE CLOUD-
BORN CHECKER-SPOT, E. nuhigena, has proven to be a delicately
marked dwarf form that has adapted itself to alpine areas. All are sub-
species of the widely ranging Etiphydryas editha Bdv., each with a well
defined and mutually exclusive range of its own.
Life Zones: Upper Sonoran, Transition for E. e. rubicunda and E. e.
monoensis; Hudsonian, Arctic-Alpine for E. e. nuhigena. Plant Com-
munities: Foothill Woodland for E. e. rubicunda; Pihon-Juniper Wood-
land for E. e. monoensis; Alpine Fell-Fields for E. e. nuhigena. Host
Plants: PLANTAGINACEAE: Plantago (Plantain).
19. Melitaea {Chlosyne) damoetas malcolmi Comst.
PL III, fig. a
MALCOLM’S CHECKER-SPOT was described from the then only
known colony at Red Lake, el. 11,000 ft., above the Mammoth Lakes
Basin, where it flies on a precipitous talus slide. It is now known to
fly in the Yosemite region and throughout the central Sierra at cor-
responding elevations. Perhaps the most intimate knowledge of the
species is possessed by Oakley Shields, who has combed the high coun-
try for isolated colonies. The most accessible of these is located above
Upper Gaylor Lake near an abandoned stone cabin reached by a fish-
erman’s trail from Tioga Pass.
Life Zones: upper Hudsonian, Arctic- Alpine. Plant Communities:
Alpine Fell-Fields. Host Plants: Not known. Possibly COMPOSITAE:
Aster ( Aster ) .
20. Melitaea {Chlosyne) acastus Ed'w.
THE ACASTUS CHECKER-SPOT is a Great Basin flier, found
sparingly around Mono Lake. The few records from late June and
early July suggest an earlier flight season.
Life Zones: Upper Sonoran. Plant Communities: Sagebrush Scrub,
Meadowland. Host Plants: COMPOSITAE: Aster (Aster).
21a. Melitaea {Chlosyne) palla palla Bdv. PL II, fig. j
b. Melitaea {Chlosyne) palla whitneyi Behr
THE NORTHERN CHECKER-SPOT ranges widely throughout
the Sierra Nevada and Cascade chains as well as the Rockies. It is re-
placed at higher elevations in the southern Sierra by WHITNEY’S
CHECKER-SPOT, a darker and ruddier form. Whether whitneyi
actually occurs in the Yosemite region is a matter for further investi-
gation. None were encountered in the course of the present survey.
Life Zones: Upper Sonoran, Transition, Canadian for M. (C.) p.
palla; Hudsonian for M. (C.) palla ivhitneyi. Plant Communities:
Foothill Woodland, Chaparral, Mixed Evergreen Forest. Host Plants:
COMPOSITAE: Aster (Aster).
2(i)ti-96, 196)
YOSEMITE BUTTERFLIESi
29
22. Melitaea {Chlosyne) hoffmanni Behr PI. II, fig. 1
HOFFMANN’S CHECKER-SPOT was the most abundant but-
terfly on the Research Reserve in 1933. They fairly swarmed over
Monardelh; the nectar seeming to intoxicate, or at least render them
oblivious to the observer’s presence. In 1956 but a few were seen on
a north-facing slope along the Tioga Road opposite the White Wolf
turnoff.
Life Zones: Canadian. Plant Communities: Openings in Fir For-
est. Host Plants: Not known.
23. Melitaea {Chlosyne) leanira F. & F.
THE LEANIRA CHECKER-SPOT invades the western portions
of the park from the foothills east of the San Joaquin Valley, where
it properly inhabits the chaparral. It is common at Indian Flat, below
El Portal.
Life Zones: Upper Sonoran (Transition). Plant Communities:
California Sagebrush Scrub (also called Coastal Sage Scrub), Foothill
Woodland. Host Plants: SCROPHULARIACEAE: Castilleja (Indian
Paint Brush), Cordylanthus tenuis (Bird’s Beak).
24a. Phyciodes {Phyciodes) campestris campestris (Behr)
b. Phyciodes {Phyciodes) campestris montana (Behr)
THE FIELD CRESCENT is found about moist places at lower
elevations, as on the floor of Yosemite Valley, or at Mather, just outside
the park. Higher up it grades imperceptibly into THE MOUNTAIN
CRESCENT, in which the tawny ground color has all but obliterated
the submarginal row of spots on the secondaries. Although heretofore
treated as a distinct species, on the basis of Yosemite specimens the
present authors prefer to consider montana but an altitudinal race of
campestris. Specimens from Mono Lake are clearly intermediate.
Life Zones: Upper Sonoran, Transition for P. (P.) c. campestris;
Canadian, lower Hudsonian for P. (P.) c. montana. Plant Communi-
ties: Several. Wet Meadowland, Roadside Associations. Host Plants:
COMPOSITAE: Aster (Aster).
25. Phyciodes {Phyciodes) mylitta (Edw.) PL II, fig. c
THE MYLITTA CRESCENT frequents the streamside and moist
meadows, often in association with the following species, Polygonia
satyrus (Edw.). It can be recognized by the bright yellow-brown
ground color. California specimens are typical, that is to say, darker,
whereas Rocky Mountain specimens incline toward the lighter races
Phyciodes ( P. ) mylitta pallida Edw. and thehais ( G. & E. ) .
Life Zones: Upper Sonoran, Transition, Canadian. Plaht Communi-
ties: Several, as is frequent with wide-ranging species. Host Plants:
COMPOSITAE: Carduus (Italian Thistle), Cirsium (Native Thistle),
and Silyhum (Milk Thistle) .
26. Polygonia satyrus { Edw. )
THE SATYR is a streamside flier that alights on the trunks of
1963
YOSEMITE BUTTERFLIES
31
Alder, Willow, and Cottonwood, to blend indistinguishably with the
mottled bark.
Life Zones: Upper Sonoran, Transition. Plant Communities: Foot-
hill Woodland, Mixed Coniferous Forest, and others. Riparian Associa-
tion. Host Plants: URTICACEAE: Urtica (Nettle).
27. Polygonia faunus rusticus ( Edw. )
THE RUSTIC ANGLE-WING occurs principally north of Yose-
mite, but may be looked for. wherever Azalea grows. Its inclusion in
the Yosemite fauna is based on an early record of Dr. E. O. Essig. The
specimen was seen by the senior author at the Yosemite Museum in
1933.
Life Zones: Transition. Plant Communities: Several. Moist Wood-
land, Riparian associations. Host Plants: ERICACEAE: Rhododendron
(Azalea); SAXIFRAGACEAE: Ribes (Currant, Gooseberry); BETU-
LACEAE: Alnus (Alder); SALICACEAE: Salix (Willow).
28. Polygonia zephyrus (Edw.) PL II, fig. k
THE ZEPHYR replaces THE SATYR {Polygonia satyrus) as the
common Angle- Wing of the Canadian and Hudsonian zones, its pale
gray underside serving as a distinguishing feature. Yosemite Creek in
mid-July is a typical situation for the species. Its food plant, Ribes^ is a
common shrub in the Subalpine Forest.
Life Zones: Transition (upper), Canadian, Hudsonian. Plant Com-
munities: Fir Forest, Lodgepole Pine Forest, Subalpine Forest. Host
Plants: ERICACEAE: Rhododendron (Azalea); SAXIFRAGACEAE:
Ribes (Currant, Gooseberry).
29. Nymphalis calif ornica {^dv.)
THE CALIFORNIA TORTOISE-SHELL appears in outbreak num-
bers every few years throughout the Sierra. The June flights observed
by the senior author in General Grant (now Sequoia-Kings Canyon)
National Park in 1922, in the San Bernardino Mountains in 1931, and
in Yosemite in 1933 were aU in a northerly or northwesterly direction,
roughly paralleling the axis of the Sierra. However, former Park
Naturalist C. A. Harwell, while on a glacier-measuring survey, reported
a remrn flight on Oaober 4, 1933, in which myriads of the inseas
passed through the highest mountain passes, all heading south. The
duration of the spring flight is but a few days. It begins at dawn and
lasts until after sundown. At times the greatest density of the insects
appears to be at tree-top level; again they come so near the ground that
they may be knocked down with a hat. A few weeks later the Ceanothus
is matted with webs of the gregarious larvae. Complete defoliation over
large areas results. The investigation of such an outbreak was the first
professional assignment of Dr. H. C. Bryant, co-founder of the Yosemite
Field School. Observations by Edwards and Behr before 1900 in the Mill
Valley seaion are mentioned by C. B. Williams in his monograph on
butterfly migration. One of us (Tilden) has observed this phenomenon
in Yosemite as recently as 1961. The explanation seems to be that the
CALIFORNIA TORTOISE-SHELL is a swarming species which, like
32
GARTH AND TILDEN
J. Ket. tepid.
the lemming, has cycles of abundance followed by a drastic reduaion
in the population, due to causes as yet not fully understood.
Life Zones: Unrestricted in flight, although the food plant does not
occur above Canadian. Host Plants: RHAMNACEAE: Ceanothus,
particularly C. cordulatus (Snow-Bush), and other smooth-leaved species.
30. Nymphalis milherti furcillata ( Say )
MILBERT’S TORTOISE-SHELL has an inward urge to fly upward,
and has been seen at the very summit of Mt. Lyell, el. 13,090 ft., the
highest point in Yosemite. Form subpdlida (CklL) flies with normal
individuals, although not commonly. The subspecific designation furcil-
lata (Say) is considered applicable to all western specimens south of
Oregon.
Life Zones: Transition, Canadian, straying higher. Host Plants:
URTICACEAE: Urtica (Nettle).
3 1 . Nymphalis antiopa ( Linn. )
THE MOURNING CLOAK is one of the world s most widespread
butterflies, occurring in Europe, northern Asia, and Japan. In England
it is called The Camberwell Beauty.
Life Zones: Unrestricted. Host Plants: SALICACEAE: Salix (Wil-
low); Populus (Poplar, Cottonwood); ULMACEAE: Ulmus (Elm).
32. V anessa atalanta (Linn.)
THE ALDERMAN or THE RED ADMIRAL is found in Europe
and Japan, as well as in North America. Few butterflies are easier to
recognize by sight.
Life Zones: Upper Sonoran, Transition. Host Plants: URTICA-
CEAE: Urtica (Nettle), Boehmeria (False Nettle), Parietaria (Pelli-
tory).-
33. V anessa virgifiiensis {T>ra,)
THE VIRGINIA LADY is widely distributed throughout North
America and may be recognized by the large eye-spots on the under
surface of the hind wings.
Life Zones: Upper Sonoran, Transition, Canadian. Host Plants:
COMPOSITAE: Gnaphalium (Cudweed; Everlasting), Anaphalis
(Everlasting, Antennaria (Everlasting), Artemesia heterophylla
(Mugwort) .
34. V anessa cardui {Unn.)
THE PAINTED LADY or THE THISTLE BUTTERFLY is prob-
ably the most cosmopolitan of butterflies. Native to the Northern
Hemisphere, it has been introduced elsewhere.
Life Zones: Unrestricted. Host Plants: BORAGINACEAE: Am-
sinckia (Fiddle-Neck), Crypantha (Nievitas); COMPOSITAE: Car-
duus (Italian Thistle), Cirsium (Native Thistle), Silybum (Milk
Thistle) .
35. Vanessa car ye {Hhn,)
THE WEST COAST LADY is restricted to the western United
2(i)ti~s6, i$6}
YOSEMITE BUTTERFLIES
33
States^ where it is more common in back yards and vacant lots than in
the out-of-doors.
Life Zones: Unrestricted. Host Plants: MALVACEAE: Malm
(Mallow), Lavatera (Tree Mallow), Sida (Alkali Mallow), sidalcea
( Checkerbloom, Wild Hollyhock), Sphaeralcea {—Malvastrum)
( Apricot Mallow, Bush Mallow ) .
36. Precis orithya evarete (Cram.)
Syn. Junonia coenia Hbn.
THE BUCKEYE may be recognized by the brightly colored eye-
spots on all wings. The aggressive males have been known to select a
perch and defend a "territory’'.
Life Zones: Upper Sonoran, Transition, (lower Canadian). Host
Plants: PLANTAGINACEAE: Plantago (Plantain) ; SCROPHULARI-
ACEAE: Mimulus (Monkey Elov/er).
37. Limenitis {Limenitis) weidemeyerii nevadae (B. & B.)
PI. IV, fig. h
THE NEVADA ADMIRAL is a Great Basin race of the Rocky
Mountain Weidemeyer’s Admiral, A (L.) weidemeyerii (Edw.),
found at Mono Lake on the eastern side of the Sierra Nevada. At Lee
Vining, where its range overlaps that of the following species, L. (L.)
lorquini Bdv., the hybrid L. (L.) fridayi (Gund.) occurs. Since the
blend is an even one, in which the characteristics of neither species pre-
dominate, we would prefer to indicate it as L. w. nevadae X T. lorquini,
rather than as a form of the above.
Life Zones: Upper Sonoran, Transition. Plant Communities: Several.
Aspen Grove, Willow Thicket, Riparian Associations. Host Plants:
SALICACEAE: Populus (Aspen, Cottonwood), Salix (Willow).
38. Limenitis {Limenitis) lorquini Bdv. PI. IL fig. b
LORQUIN’S ADMIRAL adheres to the riparian association through-
out three life zones, seldom straying from the willows and poplars of
the stream bed. On the wing it is sometimes confused with the follow-
ing species, but the manner of flight is quite different.
Life Zones: Upper Sonoran, Transition, Canadan. Plant Communi-
ties: Several. Aspen Grove, Willow Thicket and Riparian Associations.
Host Plants: SALICACEAE: Salix (Willow), Populus (Aspens and
Cottonwoods) . ROSACE AE: Prunus ( Choke -cherry ) .
39. Limenitis (Adelpha) bredowii calif ornica (Bud.)
PL I, fig. g
THE CALIFORNIA SISTER flies with the foregoing species, which
it superficially resembles, but from which it may be told on the wing by
the habit of alternating a few rapid beats with a glide in which the
wings are held slightly below the horizontal level. Unlike LORQUIN’S
ADMIRAL, however, THE CALIFORNIA SISTER forsakes the stream
margins for the hot canyon walls and the Golden Cup Oak association.
No other insect is so likely to attract the attention of the park visitor.
34
GARTH AND TILDEN
/. Res. Lepid.
Life Zones: Upper Sonoran, Transition, Canadian. Plant Communi-
ties: Oak Woodland, Mixed Evergreen Forest. Host Plants: FAGA-
CEAE: Quercus (Oak), especially Quercus chrysolepis (Golden Cup
Oak, Maul Oak ) .
40a. Apodemia mormo mormo ( F. & F. )
b. Apodemia mormo tuolumnensis Opler & Powell
THE MORMON METAL-MARK is found at higher elevations and
on the east Sierran slope. THE TUOLUMNE METAL-MARK is known
only from Pate Valley in the Grand Canyon of the Tuolumne, the type
locality. It is "a Sierran population characterized by a great extent of
orange on the upper side together with reduction of white spots and dark
under side. So far as is known it is restricted to a population along a
four mile area of the Pate Valley Trail from about 4500’ to 7000’. It is
likely a segregate from virgulti, its nearest geographic relative, which
probably occurs in scattered colonies through the lower foothills to the
west.” (Opler and Powell, 1961) BEHR’S METAL-MARK, Apodemia
mormo virgulti (Behr) , is the near relative.
Life Zones: Upper Sonoran (Transition) for A. m. tuolumnensis;
to Hudsonian for A, m. mormo. Plant Communities: Chaparral for
A. m. tuolumnensis; Sagebrush Scrub and Pi non- Juniper Woodland for
A. m. mormo. Host Plants: POLYGONACEAE: Eriogonum (Buck-
wheat) , perennial species only.
LYCAENIDAE
41. Habrodais grunus (Bdv. ) PL I, fig. h
BOISDUVAL’S HAIR-STREAK is noted for its crepuscular habits.
A party of the Yosemite Field School, on its way to the Research Reserve,
discovered it flying at Columbia Point before 5 a.m. in semi-darkness. It
continues to hover above the live oaks after the last rays of sunlight have
left the canyon walls.
Life Zones: Transition (Canadian). Plant Communities: Oak Wood-
land, Chaparral. Host Plants: FAGACEAE: Quercus chrysolepis (Gol-
den Cup Oak, Maul Oak); Castanopsis (Chinquapin) suspected.
42. Atlides halesus estesi Clench
THE GREAT BLUE HAIR-STREAK displays a metallic blue of
dazzling luster on the superior surfaces. Rarely encountered north of
Tehachapi, it has been found several times in Yosemite since the capture
by a small boy of the specimen recorded by Garth (1935). The worn
condition of Yosemite specimens suggests flight from lower elevations,
perhaps outside the park.
Life Zones : Upper Sonoran ( Transition, Canadian ) . Plant Com-
munities: Foothill Woodland. Host Plants: LORANTHACEAE :
Phoradendron ( Mistletoe ) , male flowers only.
43. Strymon {Strymon) melinus pudica (Hy. Edw.)
THE COMMON HAIR-STREAK flies in the Museum garden and
throughout the lower elevations of the park. It is the western race
pudica, the only race that flies in California.
2(i):i-96, 1963
YOSEMITE BUTTERFLIES
35
Life Zones: Upper Sonoran (Transition). Plant Communities: Sev-
eral. Host Plants: BORAGINACEAE: Cynoglossum (Hound’s
Tongue). LEGUMINOSAE: Astragalus (Loco Weed, Rattle Weed),
Hosackia {—Lotus) , Lupin us (Lupine). MALVACEAE: Malva (Mal-
low). POLYGON ACE AE: Polygonum (Knot Weed). GUTTIFERAE:
Hypericum (St. John’s Wort). LABIATAE: Hyptis (Bee Sage).
44. Strymon {Satyrium) fuliginosum semiluna (Klots)
PI. IV, fig. i
THE SEMI-LUNATE GOSSAMER WING is an early season flier
at Mono Lake. Like Melitaea {'Chlosyne) acastus, it has been found in
late June, but is probably on the wing a month earlier.
Life Zones: Upper Sonoran. Plant Communities: Sagebrush Scrub.
Host Plants: LEGUMINOSAE: Lupinus (Lupine).
45. Strymon {Satyrium) hehrii (Edw.) PL IV, fig. g
BEHR’S HAIR-STREAK is characteristic of the dry eastern slopes
of the Sierra Nevada. An isolated colony also exists on the desert slope
of the San Bernardino Mountains, in southern California..
Life Zones: Upper Sonoran. Plant Communities: Sagebrush Scrub.
Host Plants: LEGUMINOSAE: Lupinus (Lupine), Lotus {=Hos-
ackia), Astragalus (Loco Weed, Rattle Weed). ROSACEAE: Purshia
tridentata (Antelope Brush, Bitter Brush), reported (1963).
46. Strymon {Satyrium) auretorum (Bdv.) PL I, fig. k
THE GOLDEN HAIR-STREAK may now be reported authori-
tatively from the Yosemite region on the strength of specimens found
during the present survey along Hetch-Hetchy Road north of Mather.
When compared with specimens of The Nut-Brown Hair-Streak,
Strymon {Satyrium) auretorum spadix (Hy. Edw.), from Crystal Lake
in the San Gabriel Mountains of southern California, they proved to be
noticeably different in hue and marking. Specimens were attracted to
Syringa, Philadelphus Lewisii var. calif ornicus , to the fragrant blooms
of which the Theclinae are partial.
Life Zones: Upper Sonoran. Plant Communities: Chaparral. Host
Plants: FAGACEAE: Quercus (Oak), especially Q. Douglassii (Blue
Oak) in the Sierra foothills.
47. Strymon {Satyrium) saepium (Bdv.)
THE HEDGE-ROW HAIR-STREAK is abundant on the blos-
soms of the Buckeye, Aes cuius calif ornicus, in the late afternoon, where
it flies with Strymon { Satyrium ) auretorum and Euphydryas chalcedona.
Life Zones: Upper Sonoran. Plant Communities: Chaparral (west
slope); Sagebrush Scrub (east slope). Host Plants: RHAMNACEAE:
Ceanothus, especially C. cuneatus (Buck Brush). ROSACEAE: Cerco-
carpus (Mountain Mahogany) .
48. Strymon {Satyrium) adenostomatis (Hy. Edw.)
THE GRAY HAIR-STREAK occurs in the western foothills near
Jersey dale and along the road to Hetch-Hetchy, above Mather.
Life Zones: Upper Sonoran. Plant Communities: Chaparral. Host
36
GARTH AND TILDEN
/. Res. Lepid.
Plants: ROSACEAE: Cercocarpus, especially C. betuloides (Hard Tack,
Mountain Mahogany). Not Adenostoma (Chamise Bush), as its name
would imply.
49. Strymon {Satyrium) sylvinus (Bdv.)
THE SYLVAN HAIR-STREAK is found about the wiUows of the
stream bed, rather than in forest glades, as the name might suggest.
Specimens from Mono Lake show no great tendency toward race
desertorum (Grin.), which occurs from Kern County north through
Owens Valley.
Life Zones: Upper Sonoran (lower Transition). Plant Communi-
ties: Several. Riparian Association. Host Plants: SALICACEAE: Sdix
(Willow).
50. Strymqn {Satyrium) calif ornica (Edw.)
THE CALIFORNIA HAIR-STREAK is partial to the flowers of
Pink Pussy-Paws, Spraguea umbellata, from which it may be plucked
with fingers or with forceps. It is also attracted to Yerba Santa, Erio-
dictyon calif ornicum, when it visits the nature garden behind the Yose-
mite Museum.
Life Zones: Upper Sonoran (lower Transition) on west slope;
Upper Sonoran and High (arid) Transition on east slope. Plant Com-
munities: Chaparral, Oak Woodland (west slope); Sagebrush Scrub,
Juniper Woodland (east slope). Host Plants: RHAMNACEAE: Ceano-
thus cuneatus (Buck Brush). ROSACEAE: Cercocarpus (Mountain
Mahogany). Possibly FAG ACE AE: Quercus (Oak).
51. Strymon {Satyrium) dry ope (Edw.)
THE DRYOPE HAIR-STREAK was encountered playing about
sage at Mono Lake.
Life Zones: Upper Sonoran. Plant Communities: Sagebrush Scrub
(east slope) ; Chaparral, Foothill Woodland (coastal California). Ripar-
ian Association. Host Plants: SALICACEAE: Salix (Willow).
52. Callophrys {Callophrys) dumetorum (Bdv.)
THE BRAMBLE HAIR-STREAK is the green hair-streak of
moderate elevations on the western slope of the Sierra Nevada. At high
elevations, and on the eastern slope, it is replaced by the following
species.
Life Zones: Upper Sonoran. Plant Communities: Chaparral. Host
Plants: POLYGONACEAE: Eriogonum (Buckwheat), especially E.
nudum.
53. Callophrys {Callophrys) lemberti Tilden PL III, fig. k
LEMBERT’S HAIR-STREAK is the greenish hair-streak of the
higher elevations of the park. Its relationships appear to be with
Callophrys (C.) sheridani Edw. of the Rocky Mountains, in which the
band of white across the under side of the secondaries is more or less
continuous. Discovered during the course of this survey, it has been
named in honor of the pioneer resident of Tuolumne Meadows, John
Batiste Lembertr
2(i):i-96, 1963
YOSEMITE BUTTERFLIES
37
Life Zones: Hudsonian, Arctic- Alpine. Plant Communities: Alpine
Fell-Fields. Associated with "rock garden” Eriogonum, E. ovalifolium.
Host Plants: Unknown.
54. Callophrys {Mitoura) spinetorum (Hew.)
THE THICKET HAIR-STREAK is a highly prized find that may
be recognized by the bold white line across the under side of the second-
aries. Its dull bluish upper side distinguishes it from the following
Callophrys {Mitoura) johnsoni (Skin.).
Life Zones: Transition, Canadian. Plant Communities: Mixed Coni-
ferous Forest, White Fir Forest, Red Fir Forest. Host Plants: LORAN-
THACEAE: Arceuthohium (Dwarf Mistletoe), a genus occurring on
conifers.
55. Callophrys {Mitoura) johnsoni (Skin.) PL I, fig. 1
JOHNSON’S HAIR-STREAK, a close relative of Callophrys {Mi-
toura) spinetorum, may be told from the latter by means of its brownish
upper side. The species is a prize over all of its range, but occurs more
frequently in the Yuba Pass-Gold Lake area of Sierra County, California,
and in the Pacific Northwest.
Life Zones: Transition, Canadian. Plant Communities: Coniferous
Forest. Host Plants: LORANTHACEAE: Arceuthohium Douglassii
(Dwarf Pine Mistletoe) .
56. Callophrys {Mitoura) nelsoni (Bdv.) PL I, fig. i
NELSON’S HAIR-STREAK, like The California Hair-Streak,
occurs in the nature garden behind the Yosemite Museum. Both swarm
over the pink clusters of Spraguea { Pussy Paws ) and may be picked off
by forceps. They may also be found at Jerseydale, at the junction of
Highway 120 with Hetch-Hetchy Road, and on the warm road-shqulders
above Tamarack Flat.
Life Zones: Transition, Canadian. Plant Communities: Mixed Coni-
ferous Forest. Host Plants: PINACEAE: Libocedrus decurrens (Incense
Cedar) , probably.
57. Callophrys {Incisalia) doudorojfi windi (Clench)
PI. I, fig. j
WIND’S ELFIN was encountered at a damp spot in the trail near
Indian Flat, where it was flying with the following species. It differs
from normal doudoroffi of the Monterey Coast in being much lighter
and less heavily banded below. Males are slaty gray above and brown
beneath; females are quite golden in color. The form has not been
illustrated previously.
Life Zones: Upper Sonoran. Plant Communities: Chaparral. Host
Plants: Not known. Perhaps CRASSULACEAE: Sedum (Stonecrop),
as for the related C. {!.) fotis.
58. Callophrys {Incisalia) iroides (Bdv.)
THE WESTERN ELFIN is one of the first butterflies to emerge in
the spring. March finds it on the wing in the foothills of the San Joaquin
Valley. It is a very plain, brown elfin. The larvae are probably bud-
feeders, like those of Celastrina argiolus echo { Edw. ) .
38
GARTH AND TILDEN
/. Res. Lepid.
Life Zones: Upper Sonoran (lower Transition), occasionally to
Canadian. Plant Communities: Chaparral, and the chaparral-like under-
story of coniferous forests. Host Plants: CONVOLVULACEAE: Cuscuta
(Dodder). RHAMNACEAE: Ceanothus (Buck Brush, Wild Lilac).
ERICACEAE: Arhutus {M-Rdrioho) , Arctostaphylos (Manzanita), Vac-
cinium (Bilberry). Perhaps CRASSULACEAE: Sedum (Stonecrop).
59. Callophrys (Incisalia) eryphon (Bdv.)
THE WESTERN BANDED ELEIN clings quite constantly to the
coniferous forest. Never common, its discovery gives cause for comment.
Life Zones: Transition, Canadian, lower Hudsonian. Plant Communi-
ties: Yellow Pine, Red Eir, and Lodgepole Pine Forest. Host Plants:
PIN ACE AE: Pinus ponder osa (Yellow Pine), P. contorta Murray ana
(Lodgepole Pine) .
60a. Lycaena {Tharsalea) arota arota (Bdv.)
b. Lycaena {Tharsalea) arota virginiensis (Edw. )
THE AROTA COPPER and THE NEVADA COPPER occupy the
western and eastern slopes of the Sierra Nevada, respectively. The for-
mer was found in Pate Valley by Edmund Godwin on California Laurel,
Umbellularia calif ornica; the latter occurs at Mono Lake and at Bodie
where specimens seem almost indistinguishable from those from Vir-
ginia City, Nevada, the type locality.
Life Zones: Upper Sonoran, Transition. Plant Communities: Mixed
Evergreen Forest (west slope); Juniper Woodland, Sagebrush Scrub
(east slope). Host Plants: SAXIFRAGACEAE: Ribes (Currant),
Grossularia ( Gooseberry ) . The overwintering eggs are deposited at the
bases of thorns.
61. Lycaena (Lycaena) gorgon (Bdv.)
THE GORGON COPPER flies in the foothills surrounding the
western approaches to the park. Single brooded, it is on the wing during
late spring and early summer, when it may be expected to occur as a
vagrant within the park boundaries.
Life Zones: Upper Sonoran. Plant Communities: Foothill Wood-
land, Chaparral. Host Plants: POLYGONACEAE: Eriogonum (Buck-
wheat), especially E. nudum.
62. Lycaena (Lycaena) heteronea Bdv.
THE VARIED BLUE is common on the east slope of the Sierra
Nevada. It has been found west of the divide in the sagebrush zone
above the subalpine forest between Tioga Pass Checking Station and
the summit of Gay lor Lakes Trail. It is also sometimes common below
Tuolumne Meadows on the way to Return Creek, below Glen Aulin.
Life Zones: Hudsonian (in Yosemite). Plant Communities: Sage-
brush Scrub, often where mixed with Pine Forest or Juniper Woodland.
Host Plants: POLYGONACEAE: Eriogonum (Buckwheat). Asso-
ciated with E. umbellatum ( Sulphur Flower ) .
63. Lycaena (Lycaena) xanthoides (Bdv.)
THE GREAT COPPER flies in the foothills throughout the state
and is distinguished by its large size and dull coloration.
3{i):i-p6, ip6}
YOSEMITE BUTTERFLIES
39
Life Zones: Upper Sonoran (lower Transition) . Plant Communities:
Chaparral, Foothill Woodland, Yellow Pine Forest clearings. Meadow
and open Streamside. Host Plants: POLYGONACEAE: Rumex hymen-
osepalus (Dock).
64. Lycaena (Lycaena) rubidus (Behr) PI. IV, fig. d
THE RUDDY COPPER, earlier suspected of flying in the Yosemite
region (Garth, 1935), occurs in the sagebrush region of the Gay lor
Lakes Trail mentioned under the preceding Lycaena ( L, ) heteronea, and
on the east Sierran slope as well.
Life Zones: Upper Sonoran to Hudsonian (Arctic- Alpine ) . Plant
Communities: Sagebrush Scrub, openings in Subalpine Forest, and
Alpine Fell-Fields. Host Plants:. POLYGONACEAE: Rumex (Dock),
probably.
65. Lycaena (Lycaena) edit ha (Mead)
EDITH’S COPPER is a creature of the higher elevations of the
eastern portion of the park, where it flies in company with Lycaena
mariposa, L. mhidus, and L. heteronea. It resembles a compact model of
Lycaena xanthoides, and might be called ( with tongue in cheek ) , "The
Small Great Copper.”
Life Zones: Hudsonian, Arctic- Alpine. Plant Communities: Sub-
alpine Forest openings, Subalpine Meadow, Alpine Fell-Fields. Host
Plants: POLGONACEAE: Rumex paucifolius (Alpine Dock). ROS-
ACEAE: Potentilla tenuiloba, Horkelia fusca (Cinquefoil).
66. Lycaena (Lycaena) mariposa Keak.
REAKIRT’S COPPER frequents small meadows of the southern
exposure as far as timber line, but does not fly above. It was encountered
by the senior author at 10,600 feet while crossing the Lyell Fork of the
Tuolumne River preparatory to an ascent of Kuna Crest.
Life Zones: Hudsonian. Plant Communities: Openings in Subalpine
Forest. Host Plants: POLYGONACEAE: Polygonum Douglassii
(Buckwheat); can be raised on Rumex (Dock).
67. Lycaena (Lycaena) nivalis (Bdv. ) PL II, fig. f
THE NIVALIS COPPER, known for many years only from a small
colony at Glacier Point, flies generally throughout the park at elevations
of 7,000 feet and above. It resembles The Purplish Copper, Lycaena
(L.) helloides, but has an immaculate under side.
Life Zones: Canadian, Hudsonian. Plant Communities: Rocky Out-
crops and Alpine Fell-Fields. Host Plant: Unknown. POLYGON-
ACEAE: Efiogonum (Buckwheat) suspected.
68. Lycaena (Lycaena) helloides (Bdv.)
THE PURPLISH COPPER is believed to be primarily a dweller of
the lowlands, but occasional specimens have been found at higher eleva-
tions. It lacks the luster of the preceding Lycaena (L.) nivalis and has
a faintly dotted under side.
Life Zones: Upper Sonoran (Transition). Plant Communities:
Forest openings. Moist situations, rocky outcrops. Host Plants: POLY-
GONACEAE: Rumex acetosella (Sheep Sorrel), Polygonum aviculare
(Knotweed), Oxy theca. ONAGRACEAE: Gay ophy turn.
40
GARTH AND TILDEN
/. Res. Lepid.
69. Lycaena (Lycaena) phlaeas hypophlaeas (Bdv.)
PL III, fig. h
The Sierran form of THE AMERICAN COPPER, first reported
from the Yosemite region by Garth (1935b), has been established as
the typical Lycaena (L.) phlaeas hypophlaeas. Although the glacial tarn
on the north face of Mt. Maclure known locally as Bert Lake in honor
of former park naturalist C. A. "Bert” Harwell was revisited by Oakley
Shields, no additional specimens were found there. The north slope of
Mt. Dana, however, yielded a limited number, found on a steep slope
on which footing was precarious. Characterized by its small size, sub-
dued brassy color, and bluish spots above the orange subm.arginal band
of the secondaries, the Yosemite form appears closer to the Arctic race
feildeni M’Lachlan than to the eastern race, to which the name americana
Morris is now applied.
Life Zones: Arctic- Alpine. Plant Communities: Alpine Fell-Fields
Host Plants: POLYGONACEAE: Rumex (Dock), especially R. aceto-
sella, for the eastern subspecies.
70. Lycaena (Lycaena) cupreus (Edw.) PI. Ill, fig. j
THE LUSTROUS COPPER is the most intense bit of color on the
wing in Yosemite. Its fiery dress enlivens the mountain meadows. Only
Lycaena (L.) snotui (Edw.) of the Rockies, considered by Klots (1936)
and by Brown (1955) to be a subspecies of cupreus, rivals it in bril-
liance. It flies over Spraguea umbellata (—Calyptridium umbellatum)
(Pink Pussy Paws) on rock slides not far from retreating snow.
Life Zones: Hudsonian, Arctic- Alpine. Plant Communities: Alpine
Fell-Fields. Host Plants: Unknown.
71. Lepotes marina (Reak.)
THE MARINE BLUE, an Austral species, has been encountered by
us only in the Mono Lake area.
Life Zones: Lower Sonoran^ Upper Sonoran. Plant Communities:
Several over its wide range. Host Plants: LEGUMINOSAE: Medicago
(Alfalfa), Astragalus (Loco Weed, Rattle-Weed), Lathy rus (Sweet
Pea).
72. Brephidium exilis (Bdv.)
THE PYGMY BLUE, until found at Mono Lake in the final year
of this study, was recorded as present in the Yosemite region on the
strength of a single female specimen obtained by Oakley Shields at Sad-
dlebag Lake. It is the smallest of North American butterflies, with a
wing expanse of .65 in.
Life Zones: Unrestricted. Plant Communities: Alkali Sink, Salt
Marsh. Host Plants: CHENOPODIACEAE: Atriplex (Salt Bush),
Chenopodium (Lamb’s Quarters).
73. Hemiargus (Echinargus) isolus (Reak.) PL IV, fig. e
REAKIRT’S BLUE is an ubiquitous species, never common, but
occurring over a wide territory and to a considerable elevation in the
2(i):i~96, 1963
YOSEMITE BUTTERFLIES
41
Fig. 6. Moist meadow surrounded by Lodgepole Pine and Red Fir
forest. Canadian Zone. Research Reserve, el. 8,000 ft. Anthocharis sara
Stella, Plejus saepiolus, Polygonia zephyrus, Precis orythya evarete, and Pyrgus
ruralis may be seen here.^ — ^Neva Snell.
42
GARTH AND TILDEN
/. Res. Lepid.
drier eastern portions of the Sierra Nevada, at least in the 1958 season.
A row of black dots encircled with white on the underside of the pri-
maries is a distinguishing feature.
Life Zones: Unrestricted. Plant Communities: (Among others)
Sagebrush Scrub. Host Plants: LEGUMINOSAE: Astragalus (Rattle
Weed, Loco Weed), Hosackia ( = Lotus), Trifolium (Clover).
74. Lycaeides ar gyro gnomon anna (Edw. ) PL II, fig. i
THE ANNA BLUE occurs at higher elevations in the western por-
tion of the park, being found in the upper tier of timbered zones. It
is always associated with lupine.
Life Zones: Canadian, Hudsonian. Plant Communities: Fir Forest
openings. Host Plants: Undetermined. Probably LEGUMINOSAE:
Lupinus (Lupine).
75. Lycaeides melissa inyoensis Nab. PL IV, figs, i and 1
THE INYO BLUE is the name to be given to the race formerly
recognized by most Californian lepidopterists as lotis ( Lint. ) . According
to Nabokov (1949), true lotis is a race of the foregoing Lycaeides
ar gyro gnomon restricted to peat bogs of Mendocino County. THE INYO
BLUE flies in the eastern portion of the Yosemite region.
Life Zones: Unrestricted. Plant Communities: Several, including
Freshwater Marsh, Alkali Sink, Meadowland, Marsh, Cultivated Field,
Fence-Row, and Roadside Associations. Host Plants: LEGUMINOSAE:
Astragalus (Loco Weed, Rattle Weed), Hosackia { — Lotus), Lupinus
(Lupine), Medicago (Alfalfa), Glycyrrhiza lepidota (Liquorice), at
Lone Pine.
76. Plebejus {Agriades) glandon podarce (F. & F.)
PL III, fig. g
THE GRAY BLUE is perhaps the most characteristic butterfly of
the Sierran sub-alpine meadow. It is common at Tioga Pass, where at
times it fairly swarms, always in association with Dodecatheon alpinum
(Shooting Star). As soon as the sun’s rays leave the mountain valleys,
it settles upon the grasses and sedges and may be picked by hand or by
forceps. A cross on each fore-wing serves to distinguish the species.
Life Zones: Upper Canadian, Hudsonian (Arctic- Alpine ) . Plant
Communities: Sub-alpine Meadow, Alpine Fell-Fields. Host Plants:
Unknown. PRIMULACEAE: Gregoria for European P. glandon, to
which Primula (Primrose) and Dodecatheon (Shooting Star) are re-
lated.
77. Plebejus {Plebejus) saepiolus (Bdv.)
THE GREENISH BLUE flies with the foregoing species, Plebejus
(A.) glandon podarce, and in equal abundance. P. (P.) saepiolus, how-
ever, descends to lower elevations, as at Glacier Point and Aspen Valley.
As with the former species, only the male wears the color indicated by
the common name, the female being drab brown. The commonest blue
in the park at higher elevations, and the one with the longest flight
season, it has two broods, the first tending to oviposit on clover, the
second on Hosackia.
2(i);i-96, 1963
YOSEMITE BUTTERFLIES
43
Life Zones: Canadian, Hudsonian (Arctic- Alpine). Plant Com-
munities: Subalpine Meadow, Alpine Fell-Fields, Fiost Plants: LEGUM-
INOSAE: Trifolium (Clover), Hosackia (= Lotus).
78a. Pie be jus {Icaricia) icarioides icarioides (Bdv.)
b. Plebejus {Icaricia) icarioides helios (Edw. )
BOISDUVAL’S BLUE is a widely distributed species, having been
subdivided into numerous geographical races. The Yosemite form is
close to the nominate form; the name helios Edw. is here applied to spec-
imens from Mono Lake. Because of its relatively large size and uniform
markings, as well as its solid blue coloration, this species may be confused
with Lycaena (L.) heteronea Bdv. when on the wing.
Life Zones: (Upper Sonoran), Transition, Canadian. Plant Com-
munities: Coastal Sage Scrub, Chaparral, Foothill Woodland, Yellow
Pine and Mixed Coniferous Forest openings for P. i. icarioides; Sage-
brush Scrub, Pinon-Juniper Woodland, Juniper Woodland, Jeffrey Pine
Forest openings for P. i. helios. Flost Plants: LEGUMINOSAE: Lfipinus
(Lupine) , perennial species only.
79. Plebejus (Icaricia) shasta comstocki (Fox) PL III, fig. e
THE YOSEMITE BLUE is the name given this insect in Comstock’s
"Butterflies of California,” although by virtue of its Latin name it might
be called Comstock’s Blue instead. Its metropolis is above timber line,
but isolated colonies occur above 7,000 feet, as at Glacier Point. It flies
over dwarf lupine and yellow buckwheat, Eriogonum incanum, but has
not been observed to oviposit on either. The long flight season suggests
two broods. The Rocky Mountain Plebejus (Icaricia) shasta minnehaha
(Scud.) flies only above tree line and often above 12,000 feet.
Life Zones: Hudsonian, Arctic- Alpine. Plant Communities: Sub-
alpine Forest openings, Alpine Fell-Fields. Host Plants: Unknown. LE-
GUMINOSAE: Lupinus (Alpine Lupine) suspected.
80. Plebejus ( Icaricia ) acmon ( West. & Hew. )
THE ACMON BLUE may be found throughout the Sierra Nevada
at almost any elevation. It occasionally flies in company with the follow-
ing Plebejus (I.) lupini. From this circumstance, plus a possible differ-
ent host plant, it has been suggested that the two are distinct species, and
they are tentatively so considered here.
Life Zones: Upper Sonoran, Transition, Canadian (Hudsonian. Plant
Communities: Many. Host Plants: LEGUMINOSAE: Astragalus (Loco
Weed, Rattle Weed), Hosackia ( — Lotus) , Lupinus (Lupine) . POLY-
GON ACE AE : Eriogonum ( Buckwheat ) .
81. Plebejus (Icaricia) lupini (Bdv.)
THE LUPINE BLUE may be distinguished from Plebejus (I.)
acmon by the purplish luster of both sexes. The two are sometimes
found together, as at Smoky Jack Camp. Specimens from 11,000 feet
are visibly smaller than those from moderate elevations; those from Lee
Vining are visibly larger.
Life Zones: Canadian, Hudsonian, High Arid Transition. Plant
Communities: Not determined. Subalpine Forest, Alpine Fell-Fields,
44
GARTH AND TILDEN
/. Res. Lcpid.
Juniper Woodland, Sagebrush Scrub, among others. Host Plants: Ap-
parently unknown, as distinct from P. (I.) acmon. Eriogonum (Buck-
wheat) association noted.
82. Everes comyntas (Godt.)
THE EASTERN TAILED BLUE is the species represented on the
east side of the Sierra Nevada, although it is not known to which of
several subspecies the Mono Lake population should be assigned.
Life Zones: Upper Sonoran (Hudsonian). Plant Communities: Not
determined. Freshwater Marsh, Moist Meadow associations. Host Plants:
LEGUMINOSAE: Trifolium (Clover), Vicia (Vetch), Astragalus
(Loco Weed, Rattle Weed).
83. Everes amyntula
THE WESTERN TAILED BLUE has been seen in Yosemite Park
by one of us ( Garth ) . It was flying over a tall, succulent lupine in the
vicinity of a white fir grove at Hetch-Hetchy summit. It occurs in the
foothills west of the park.
Life Zones: Transition (Canadian). Plant Communities: Yellow
Pine Forest openings. Host Plants: LEGUMINOSAE: Astragalus (Loco
Weed, Rattle Weed), Lathyrus (Sweet Pea), Vicia (Vetch).
84. Philotes enoptes ( Bdv. )
THE DOTTED BLUE, as found at Glacier Point, is typical. Speci-
mens found at Research Reserve show mixed characters, but are separable
from the following species.
Life Zones: Transition, Canadian (Hudsonian). Plant Communities:
Understory of several. Host Plants: POLYGONACEAE: Eriogonum
(Buckwheat) of the latifolium group. The eggs are concealed in the
flower head, on which the larvae feed.
85a. Philotes hattoides hattoides (Behr)
b. Philotes hattoides glaucon ( Edw. )
THE SQUARE-SPOTTED BLUE of Yosemite is an inhabitant of
the Alpine Fell-Fields, appearing very early in the year for so high an
altitude. It also flies atop Research Ridge (Boundary Hill), keeping
well within the few acres marked by the presence of weak Hudsonian
elements. It flies over Eriogonum incanum, a yellow buckwheat, and
Astragalus bolanderi, a straggly legume. Specimens found within the
park represent the typical form. Specimens found at Mono Lake are
referred to THE GLAUCON BLUE, a subspecies.
Life Zones: Hudsonian for P. h. hattoides; Upper Sonoran, Arid
Transition for P. b. glaucon. Plant Communities: Rock Garden for P. h.
hattoides; Juniper Woodland, Sagebrush Scrub for P. b. glaucon. Host
Plants: POLYGONACEAE: Eriogonum (Buckwheat).
86. Philotes speciosa ( Hy. Edw. )
THE SMALL BLUE is typically a desert insect, found around dry
lake beds (playas) in the Mojave Desert. Occasional specimens occur
in the Central Valley region of California, and one of us (Garth) has
found it at Hmne Lake in Sequoia-Kings Canyon National Park. The
2(t):i-96, 1963
YOSEMITE BUTTERFLIES
45
Mariposa and Briceburg specimens collected by G. and R. Bohart may
represent an undescribed subspecies. Both specimens are larger, with
wider margins in the male, and with the hind wings less spotted below,
than normal specimens from the desert. In the absence of sufficient
material, description must wait.
Life Zones: Lower Sonoran in the Mojave Desert; Upper Sonoran
(Transition) in the western Sierra Nevada. Plant Communities: Not
determined. Stream Bed Association. Host Plants: POLYGONACEAE:
Eriogonum (Buckwheat) in San Diego County; Oxy theca perfoliata in
the Mojave Desert; in the Sierra unknown.
87» Phaedrotes piasus i'B&v.)
THE ARROWHEAD BLUE, while not common in the Yosemite
region, occurs in the western foothills, the middle elevations, and on the
eastern slope at Mono Lake.
Life Zones: Upper Sonoran, Transition, Canadian. Plant Commun-
ities: Several. Host Plants: LEGUMINOSAE: Lupinus (Lupine).
88a. Glaucopsyche lygdamus hehrii ( Edw^. )
b. Glaucopsyche lygdamus Columbia (Skin.)
BEHR’S BLUE flies in the early spring in the western foothills.
At higher elevations it is replaced by THE COLUMBIA BLUE, a
montane subspecies found in cooler parts of most of the United States
and Canada. It is not common in the Tioga Pass region, nor indeed
anywhere, its colonies being widely scattered.
Life Zones: Upper Sonoran for G. 1. hehrii; Canadian, Hudsonian
for G. 1. Columbia. Plant Communities: Foothill Woodland, Chaparral
for G. 1. hehrii; Subalpine Forest, Alpine Fell-Fields for G. /. Columbia.
Host Plants: LEGUMINOSAE: Lottis (= H os ac kia) , Astragalus (Loco
Weed, Rattle Weed), Lathyrus (Sweet Pea), Lupinus (Lupine), Vida
(Vetch).
89. Celastrina argiolus echo (Edv^. )
THE ECHO BLUE is the western representative of a highly poly-
morphic species, now recognized as having Old World affinities. One
of the earliest butterflies to emerge in the spring, it follows the season’s
advance to higher altitudes, occurring through three life zones. Male
and female are strikingly dissimilar in pattern.
Life Zones: Upper Sonoran, Transition, Canadian. Plant Commun-
ities: Many. Host Plants: LEGUMINOSAE: Hosackia (= Lotus),
Lupinus (Lupine); RHAMNACEAE: Ceanothus (Lilac), Spiraea
(Spirea); SAPINDACEAE: Aesculus (Buckeye); ERICACEAE:
Arctostaphylos (Manzanita); FAGACEAE: Quercus (Oak); CORN-
ACEAE: Cornus (Dogwood). In many instances, only the flower buds
are fed upon.
PLATE I
3(i):i-96, 1963
YOSEMITE BUTTERFLIES
47
PLATE I
THE WESTERN FOOTHILLS
a. Thorbes diversus
Aspen Valley, J.S.G.
b. Argynnis z. zerene cf
Mather, J.S.G.
c. Coenonympha t. calif ornica cf
Jerseydale, J.S.G.
d. Euphydryas e. rubicunda d
Indian Flat, J.W.T.
e. Ce rcyonis sylvestris 9
Hetch-Hetchy, J.S.G.
f. Anthocharis lanceolata d
Yosemite Valley, Paul’ Allen
g. Limenitis b. californica 9
Hetch-Hetchy, J.S.G.
h. Habrodais grunus 9
Hetch-Hetchy, J.S.G.
i. Callophrys nelsoni 9
Hetch-Hetchy, J. S.G.
j. Callophrys d. windi 9
Indian Flat, J.W. t.
k. Strymon auretorum 9
Hetch-Hetchy, J S.G.
l. Callophrys johnsoni d
Jerseydale, A.O.S.
PLATE m
THE HIGH COUNTRY
a. Melitaea d. malcolmi d
Gaylor Lakes, J.S.G..
b. Oeneis c. ivallda d
Glacier Lodge, J.S.G.
c. Euphydryas e. nubigena 9
Rafferty Creek, J.S.G.
d. Polites s. tecunaseh
Upper Gaylor Lake, J.S.G.
e. Plebejus s. comstocki d
Yosemite Park, J.S.G.
f. Parnass|us p. behri d
Gayl Lakes, J.S.G.
g. Plebejus g. podarce d
Research Reserve, J.S.G.
h. Lycaena p. hypophlaeas 9
N. slope, Mt. Dana, A.O.S.
i. Colias behri d
Tioga Pass, J.S.G.
j. Lycaena cupreus d
Lembert Dome, J.S.G.
k. Callophrys lemberti (paratype) d
Tioga Pass, J.W.T.
l. Argynnis m. arge 9
Tioga Pass, J.S.G.
m. Hesperia miriamae 9
Unicorn Peak, A.O.S.
PLATE II
THE WESTERN MID- ELEVATIONS
a. Anthocharis s. Stella 9
White Wolf, J.S.G.
b. Limenitis lorquini d
Aspen Valley, J.S.G.
c. Phyciodes mylitta 9
YosemiLe Creek, J.S.G.
d. Boloria epithore 9
Tioga Road, J.S.G.
e. Parnassius c. baldur d
White Wolf, J.S.G.
f. Lycaena nivalis 9
Research Reserve, J.S.G.
g. Hesperia harpalus cT
Yosemite Creek, J.S.G.
h. Argynnis a. irene rf
Aspen Valley, J.S.G.
i. Lycaeides a. anna cf
Aspen Valley, J.S.G.
j. Melitaea palla d
White Wolf, J.S.G.
k. Polygonia zephyrus 9
Tuolumne Grove, J.S.G.
l. Melitaea hoffmanni 9
White Wolf, J.S.G.
PLATE IV
THE MONO BASIN
a. Coenonympha t. mon o d
W of LeeVining, A.O.S.
b. Argynnis c. nevadensis d
near Mammoth, J.S.G.
c. Cercyonis oeta d
June Lake, J.S.G.
d. Lycaena rubidus d
Virginia Lakes, J.S.G.
e. Hemiargus isolus 9
Rafferty Cr., J.S.G.
f. Euphydryas e. monoensis d
Mono Lake, A.O.S.
g. Strymon behrii 9
Mammoth, J.S.G.
h. Limenitis w. nevadae d
Mono Lake, J.S.G.
i. Lycaeides m. inyoensis ff
Mono Lake, J.W.T.
j. Strymon f. semiluna
Mono Lake, A.O.S.
k. Argynnis n. apacheana (f
Round Valley, J.S.G.
l. Lycaeides m. inyoensis 9
Lone Pine, J.W.T.
PLATE II
PLATE III
50
GARTH AND TILDEN
/. Res. Lepid.
PIERIDAE
90a, Anthocharis {Anthocharis) sara saral^wc^is
Anthocharis {Anthocharis) sara reakirtii Edw., gen. vern.
b. Anthocharis {Anthocharis) sara Stella Edw. PI. II, fig. a
THE SARA ORANGE-TIP and its overwintering form, REA-
KIRT’S ORANGE-TIP, occur in the western lowlands at Briceburg and
El Portal. THE STELLAR ORANGE-TIP is the high-altitude race
found in California, race jzdia Edw. being from Colorado and race flora
Wgt. from Oregon and north. Females of this form are yellowed, un-
like those of the lowland forms, in which females are white like the
males.
Life Zones: Upper Sonoran, lower Transition for A. sara sara; upper
Transition, Canadian for A. s. Stella. Plant Communities: Valley Grass-
lands, Foothill Woodland, Chaparral, Yellow Pine Forest for A. s. sara;
White Fir Forest, Red Fir Forest for A. s. Stella. Forest openings and
Riparian Association. Host Plants: CRUCIFER AE: Arabis (Rock
Cress), Brassica (Mustard), Sisymbrium (Hedge Mustard).
91. Anthocharis {Falcapica) lanceolata Lucas PI. I, fig. f
THE LANCEOLATE MARBLE (formerly called Boisduvals
Marble when authorship was attributed to him) is an early flier, to be
sought from mid-May to mid- June. It seeks the vertical walls of Yose-
mite Valley and may be seen on the precipitous talus slopes near Bridal
Veil Falls. Yosemite specimens are almost gray enough beneath to match
race australis { Grin. ) from southern California.
Life Zones: Upper Sonoran, Transition. Plant Communities: Chap-
arral, Foothill Woodland, Mixed Evergreen Forest. Rocky Outcrop and
Cliffside Associations. Host Plants: CRUCIFER AE, especially Arabis
perfoliata (Rock Cress).
92. Euchloe creusa hy antis (Edw.)
EDWARDS’ MARBLE may be separated from the following Colo-
rado Marble by the heavy green marbling of the under side of the
secondaries against a background of dead white, Euchloe ausonides
coloradensis having less green and of a decidedly yellowish cast against
a pearly white ground color. The green of both species is an effect
produced by yellow scales laid over black, as examination with a hand
lens will demonstrate. The black bar at the end of the cell is wide in
creusa.
Life Zones: Upper Sonoran, Transition, Canadian. Plant Commun-
ities: Chaparral, Foothill Woodland, Mixed Evergreen Forest, White Fir
Forest. Host Plants: CRUCIFERAE: Arabis (Rock Cress), Streptan-
thus (Jewel Flower).
93. F.uchloe ausonides coloradensis (Hy. Edw.)
THE COLORADO MARBLE is the common Euchloe of the Re-
search Reserve, where approximately four specimens were found for
every one of E. creusa hyantis. Both fly to the mountain tops and may
be seen in good numbers by the observer who stations himself in such
PLATE IV
52
GARTH AND TILDEN
/. Res. Lepid.
a situation. Like Behr’s Parnassian, Parnassius phoebus behrii Edw., it
belongs to the Great Basin fauna. The black bar at the end of the cell
is narrow.
Life Zones: Canadian, Hudsonian. Plant Communities: Coniferous
Forest openings. Host Plants: CRUCIFER AE: Arabis (Rock Cress),
Erysimum (Wall Flower), Sisymbrium (Hedge Mustard).
94. CoUas eury theme Bdv.
THE CLOUDED SULPHUR is partial to grassy meadows and to
cultivated areas, where it substitutes introduced alfalfa for indigenous
Astragalus for its food plant. The summer form, amphidusa Bdv., is
larger and more orange. Both have two forms of the female, normal
and albinic, but only One form of the male.
Life Zones: Unrestricted. Plant Communities: Immaterial, if
legumes be present. Host Plants: LEGUMINOSAE: Astragalus (Loco
Weed, Rattle Weed), Hosackia (— Lotus) ^ Medicago (Alfalfa), Mel-
ilotus (Sweet Clover), Trifolium (Clover), Vida (Vetch).
95. CoUas philodice hagenii Edw.
HAGEN’S SULPHUR is a Great Basin race of a widely ranging
North American species found otherwise mostly east of the Rocky
Mountains. According to Hovanitz (1943, 1950), it does not occur
west of the Sierra-Cascade Divide Triple-brooded at Mono Lake, it
may be recognized by its yellow, rather than orange, coloration.
Life Zones: Upper Sonoran, Transition. Plant Communities: Moist
Meadows. Host Plants: LEGUMINOSAE: Trifolium (Clover).
96. CoUas occidentaUs chrysomelas Hy. Edw.
THE GOLDEN SULPHUR is the largest member of its genus
occurring in California. An early flier, it is considered a rarity, having
been seen by but one of us (Tilden) in the vicinity of Mather on June
30, 1962, in the course of this survey.
Life Zones: Transition. Plant Communities: Associated with con-
iferous forest at moderate elevations. Host Plants: LEGUMINOSAE:
( Legumes ) .
97. CoUas behrii Edw. PI. Ill, fig. i
BEHR’S SULPHUR is the only greenish sulphur butterfly of the
Sierra Nevada. Until the opening of the Tioga Road in 1915 it was
considered a great rarity because of the extreme inaccessibility of its
habitat, the Tuolumne Meadows. John Batiste Lembert, who homestead-
ed the Soda Springs quarter section in 1885, found that there was a
demand for these butterflies and for a dozen years supplied them in
quantity to universities and museums. The secret of their haunts he
shared with the Indians, who in turn refused to divulge it to outsiders.
We now know that CoUas behrii has as its metropolis the Tuolumne
Meadows, that smaller colonies are present in adjacent subalpine
meadows, and that it has been found to the south at Rock Creek and
Mineral King. Its occurrence is tied closely to the distribution of
Dwarf Bilberry, on which the larvae exclusively feed. Form 9 canescens
i$6s
YOSEMITE BUTTERFLIES
53
PLATE V
a. Erynnis pacuvius lilias d
Chowchilla Mtn. J.W.T.
b. Argynnis zerene malcolmi d
Mammoth Lakes, J.W. Friday
c. Lycaena gorgon 9
Lebec, J.S.G.
d. Everes amyntula d
Yucaipa, J.S.G.
e. Papilio indra d
Tioga Pass, A.O.S.
f. Callophrys iroides d
Grant Nat'l Park J.S.G.
g. Strymon saepium 9
Hetch-Hetchy, J.S.G.
h. Lycaena editha 9
Tioga Pass, J.S.G.
i. Philotes battoides glaucon d
Mono Lake. A.O.S.
j. Brephidium exilis
Long Beach, J.S.G.
k. Philotes battoides battoides d
Research Reserve, J.S.G.
l. Pieris occidentalis calyce 9
W of Tioga Pass, A.O.S.
m. Danaus plexippus d
Mather, J.S.G.
n. Euchloe creusa lotta 9
Darwin, R.P. Allen
o. Polites sonora 9
Aspen Valley, J.S.G.
p. Vanessa carye d
Topanga Canyon, J.S.G.
q. Vanessa cardui 9
Grant Nat'l Park, J.S.G.
r. Polites sonora d
Bridal Veil Creek, J.S.G.
s. Lycaena heteronea 9
Gem Lake, J.S.G.
t. Callophrys iroides d
Mammoth Lakes, J.S.G.
u. Phaedrotes piasus d
Tujunga, J.S.G.
V. Plebejus icarioides helios 9
Warren Creek, J.S.G.
PLATE VI
a . Thorbes pylades j
Jersydale, J.W.T.
b. Pyrgus ruralis cf
Boulder Creek, J.W.T.
c. Ochlodes agricola d
Hetch-Hetchy, J.S.G.
d. Amblyscirtes vialis , d
Flagstaff, Ariz., K. Roever
e. Argynnis atlantis irene d
Aspen Valley, J.S.G,
f. Leptotes marina 9
Long Beach, J.S.G.
g. Vanessa virginiensis 9
Santa CruzMtns., J.S.G,
h. Strymdn melinus pudica d
Long Beach, J.S.G.
i. Nymphalis milberti 9
Yucaipa, J.S.G.
j. Lycaena arota virginiensis 9
Mono Lake, A.O.S.
k. Pieris napi venosa d
Briceburg, K. T.
l. Papilio eurymedon ^
Idyllwild, J.S.G.
m. Plebejus saepiolus (f
Research Reserve, J.S.G.
n. Colias eurytheme cf
Mather, J.S.G.
o. Argynnis cybele leto 9
Strawberry, J.S.G.
p. Argynnis callippe inornata 9
Jerseydale, A.O.S.
54
GARTH AND TILDEN
Lepid.
J. Res.
PLATE V
YOSEMITE BUTTERFLIES
55
s(i):i-$6) 1963
56
GARTH AND TILDEN
/. Res. Lepid.
Comst., although relegated to synonymy by McDunnough (1938), we
find to be a valid albinic form that may be recognized on the wing.
Life Zones: Hudsonian (Arctic- Alpine ) . Plant Communities:
Subalpine Meadows; basin areas of Subalpine Forest; moister parts of
Alpine Fell-Fields. Host Plants: ERICACEAE: Vaccinium caespitosum
(Dwarf Bilberry).
98. Colias {Zerene) eurydice^dY.
THE CALIFORNIA DOG-FACE or FLYING PANSY, as it is
sometimes called, must be a vagrant in the Yosemite region, for Hall
(1912) does not record its food plant, False Indigo or Lead Bush, as
occurring in the park. It is abundant in the foothills of southern Cali-
fornia, and has been selected as the State butterfly. The male bears the
figure of a dogs head on the forewing; the female is pure sulphur
yellow.
Life Zones: Upper Sonoran, lower Transition. Plant Communities:
Not determined for the Yosemite region. Host Plants: LEGUMI-
NOSAE: Amorpha calif ornica (False Indigo).
99. Nathalis iole Bdv.
THE DAINTY YELLOW is likely to be encountered in the
least expected places, from sea level to 9,000 feet, and from early
spring to early winter. There are no set rules of behavior for this
diminutive sulphur, which has followed the Filaree into many an
improbable situation.
Life Zones: Unrestricted. Plant Communities: Ubiquitous. Host
Plants: GERANIACEAE: Erodmm (Filaree) . COMPOSITAE: Helen-
ium ( Sneezeweed ) , Dyssodia (None). CARYOPHYLLACEAE: Stel-
laria ( Chickweed ) .
100. Neophasia menapia tau (Scud.)
THE PINE WHITE reaches the peak of its flight in August.
The adults are high fliers, seldom descending from the tree tops. Al-
though not abundant in 1933, they were seen by the hundreds in
1928 along the Big Oak Flat road by one of us (Garth) . The larvae are
of special interest to forest entomologists, having been known to
defoliate pine trees over large areas. Resurrected from synonymy, the
name tau is now applied to the western form.
Life Zones: Transition, Canadian. Plant Communities: Yellow
Pine Forest, Jeffrey Pine Forest (east slope); occasionally Lodgepole
Pine Forest. Host Plants: PINACEAE: Pinus ponder osa (Yellow
Pine), P. contorta Murray ana (Lodgepole Pine); also, according to
the observation of one of us (Tilden), Pseudotsuga taxifolia (Douglas
101. Pieris (Pontia) beckeriiEdw.
BECKER’S WHITE prefers an arid environment and is encountered
east of the Sierra Nevada from the Mojave Desert north through
Owens Valley to Mono Lake. The green-margined veins of the under
side suggest one of the Marbles, and where it flies with Anthocharis
YOSEMITE BUTTERFLIES
57
PLATE Vn
a. Epargyreus clarus 9
Lagunitas, E.C. VanDyke
b. Hesperia juba tf
Mather, J.W.T.
c. Ochlodes sylvanoides d*
Virginia Lakes, J.S.G.
d. Erynnis propertius d
Yosemite Creek, J.S.G.
e. Glaucopsyche lygdamus Columbia d
White Wolf, J.S.G.
f. Cercyonis pegala ariane 9
Round Valley, J.S.G.
g. Argynnis egleis d
Yosemite Creek, J.S.G.
h. Plebejus i. icarioides 9
Aspen Valley, J.S.G.
i. Euchloe creusa lotta 9
Mono Lake, A.O.S.
j. Philotes speciosa d
Mojave Desert, J.S.G.
k. Heliopetes ericetorum d
Switzer-land, J.S.G.
l. Apodemia m. mormo
Mono Lake, A.O.S.
m. Papilio rutulus d
Idyllwild, J.S.G,
n. Precis orithya evarete d
Grant Nat'l Park, J.S.G,
o. Plebejus lupini
W, of Tioga Pass, A.O.S.
p. Nymphalis californica 9
Idyllwild, J.S.G.
q. Colias eurytheme form alba 9
Big Bear Lake, J.S.G.
r. Lycaenopsis argiolus echo 9
Yucaipa, J.S.G. •
s. Lycaena a. arota 9
Gold Creek, J.W.T.
t. Lycaena xanthoides 9
Mather, J.S.G.
u. Callophrys spinetorum d
Idyllwild, J.S.G.
V. Strymon adenostematis 9
Hetch-Hetchy, J.S.G.
PLATE Vm
a* Erynnis tristis d
Redwood City, J..W.T.
b. Thorbes mexicana nevada d
Tioga Pass, A.O.S.
c. Pyrgus communis d
Los Angeles, J.S.G.
d. Nathalis iole d
Warren Creek, A.O.S.
e. Pieris sisymbrii d
Research Reserve, J.S.G.
f. Ochlodes agricola d
El Portal
g. Strymon dr yop e (f
Mono Lake. A.O.S.
h. Papilio zelicaon 9
Mather, J.S.G.
i. Strymon californica d
Hetch-Hetchy
j. Lycaena helloides d
Mather, J.S.G.
k. Lycaena nivalis 9
Glacier Point, J.D. Gunder
l. Melitaea leanira d
m. Colias (Zerene) eurydiced
Santa AnaR,, J.S.G,
n. Phyciodes c. campestris d
Mather , J. S. G.
o. Argynnis mormonia arge d
Hetch-Hetchy, J.S.G.
p. Plebejus acmon d
Hatch- Hetchy
q. Philotes enoptes 9
Research Reserve, J.S.G.
r. Polygonia satyrus d
Mono Lake, A.O.S,
/. Re$. Lepid.
58
GARTH AND TILDEN
PLATE VII
V
4.
2(i):i-96, 196}
YOSEMITE BUTTERFLIES
59
PLATE
60
GARTH AND TILDEN
/. Res. Lepid.
lanceolata australis Grin, in some of the desert canyons the two are
not always distinguishable when on the wing.
Life Zones: Upper Sonoran, Transition. Plant Communities: Sage-
brush Scrub, Shadscale Scrub, Pihon-Juniper and Juniper Woodland.
Host Plants: CAPPARIDACEAE: Isomeris arbor ea (Bladder Pod).
CRUCIFER AE: Brassica nigra (Mustard), Sisymbrium altissimum
( Hedge Mustard ) , Stanley a pinnata ( Desert Plume, Prince s Plume ) .
102. Pieris {Pontia) sisymbriiBd\.
THE CALIFORNIA WHITE is an early flier in the western
foothills, where it may be found at Briceburg and El Portal in mid-
March. At higher elevations it flies in mid-summer with the follow-
ing Pieris {Pontia) occidentalis calyce, these two being the only whites
then on the wing. While Sisymbrium, from which it takes its scientific
name, grows abundantly in the meadows of Research Reserve, the
species exhibits a marked preference for Caulanthus and Streptanthus
when these are available, according to Hovanitz (personal communica-
tion).
Life Zones: Unlimited. Plant Communities: The entire gamut from
west to east. Host Plants: CRUCIFER AE: Arabis (Rock Cress),
Caulanthus (Rock Cabbage), Sisymbrium (Hedge Mustard), Strep-
tanthus (Jewel Flower).
103. Pieris {Pontia) protodice Bdv. & Lee.
THE COMMON WHITE appears limited to the lower tier of
life zones in Yosemite. Those seen above Transition Zone are believed
to have wandered upward because of their worn condition.
Life Zones: Upper Sonoran, Transition (Canadian). Plant Com-
munities: Many. Host Plants: CAPPARIDACEAE: Cleome lutea (Bee
Plant). CRUCIFERAE: Brassica nigra (Mustrad), Raphanus (Rad-
ish), Lepidi'um densiflorum (Pepper Grass), Sisymbrium altissimum
(Hedge Mustard), the first and two last at Mono Lake on the authority
of W. Hovanitz.
104a. Pieris {Pontia) occidentalis occidentalis Reak.
b. Pieris {Pontia) occidentalis calyce Edw^.
THE WESTERN WHITE occurs throughout the park, its altim-
dinal form, EDWARD’S WHITE, at higher elevations only, although
a small, dark form resembling P. o. calyce may be found in early
spring in the western foothills. What one finds in the high country
in August is calyce, freshly emerged (It is the only brood at higher
elevations), plus worn second-brood P. o. occidentalis that have in-
vaded from lower levels. L. G. Higgins of England recently (1953)
associated calyce Edw. with the European callidice Esper as subspecies
of P. {Pontia) occidenalis, which has been shown by Hovanitz (1963)
to be specifically distinct from the preceding P, {Pontia) protodice.
Life Zones: Unrestricted for P. o. occidentalis; Canadian, Hudsonian
for P. 0. calyce. Plant Communities: Many for P, o. occidentalis; Alpine
YOSEMITE BUTTERFLIES
61
Fell-Fieids for P. o. calyce. Host Plants: CRUCIFERAE: Arabis (Rock
Cress), Lepidium densijlorum (Pepper Grass), Sisymbrium altissimum
(Hedge Mustard), the last two in the Mono Lake area, according to
Hovanitz (personal communication).
105. Pieris {Pieris) napi venosa Scud,
THE VEINED WHITE is found in the foothills adjacent to Yose-
mite National Park on the San Joaquin Valley side from mid-March
to mid-May, when it appears on the floor of Yosemite Valley. More
lightly marked than typical venosa, this foothill population is not
referable to any other named subspecies.
Life Zones: Upper Sonoran, Transition. Plant Communities: Foot-
hill Woodland, Chaparral, Mixed Coniferous Forest. Riparian and
Cool Woodland associations. Host Plants: CRUCIFERAE: Barb area
(Winter Cress), Brassica (Mustard), Dentaria (Milk Maids), Rapha-
nus ( Radish ) .
106. Pieris (Pieris) rapae (Linn.)
THE CABBAGE WHITE was introduced from Europe into Que-
bec, Canada, in 1858, reaching California about 1883. It now occupies*
not only cultivated areas, but has expanded into wild lands as well.
Life Zones: Unrestricted. Plant Communities: Many, as would be
expected of a highly adaptable immigrant. Host Plants: CRUCIFERAE:
Brassica (Mustard), Radicula (Yellow Cress), the latter around
Cathay.
PAPILIONIDAE
107. Papilio zelicaon Luc.
THE ANISE SWALLOWTAIL is a "hill-topper” species, choosing
the most exposed situations for its flight. Usually only the males disport
themselves in this way, six being found at the very summit of Research
Ridge. It is partial to the blossoms of the Western Wallflower, Erysi-
mum asperum. Although readily confused with Papilio bairdii brucei
Edw., which occurs in the region of the Mono Craters east of the Sierra
Nevada, P. zelicaon has less yellow on the abdomen than does P. brucei.
Life Zones: Unrestricted. Plant Communities: Chaparral, Foothill
Woodland, openings in Coniferous Forest. Host Plants: UMBEL-
LIFER AE: Daucus (Wild Carrot), Poeniculum vulgar e (Fennel, An-
ise), Heracleum lanatum (Cow Parsnip).
108. Papilio indr a Reak.
THE INDRA SWALLOWTAIL has been seen also at the summit
of Research Ridge. An erratic flier, it has proven most elusive, and
Yosemite records are few. These include the floor of Yosemite Valley,
the summit of Yosemite Falls, northwest above Tioga Pass, and Mono
Lake. On the west bank of Yosemite Creek it flies with Parnassius
phoebus behrii Edw. on dry slopes in association with Juniper, Sedum,
and Sanicula.
62
GARTH AND TILDEN
/. Re$. Lepid.
Life Zones: Transition, Canadian, Hudsonian. Plant Communities:
Juniper Woodland. Host Plants: UMBELLIFERAE: Pteryxia ( =
Cymoptems) terebrinthina in the Sierra Nevada (Emmel and Emmel,
1963).
109. Papilio rutulus Luc.
THE WESTERN TIGER SWALLOWTAIL is a species that
transcends zonal limits in its adherence to the stream-bank associa-
tion. It occurs at El Portal on the Merced River, thence through the
Yosemite Valley and its tributary streams well into Canadian Zone.
One of us (Tilden) has even observed it in Subalpine Forest at Tioga
Pass.
Life Zones: Unrestricted. Plant Communities: Willow, Cotton-
wood, and Aspen groves. Riparian Association. Host Plants: SALI-
CACEAE: Populus (Cottonwood), Salix (Willow). BETULA-
CEAE: Alnus (Alder). ROSACEAE: Prunus (Choke-Cherry).
110. Papilio multicaudatus Kby.
Syn. Papilio daunus Bdv.
THE TWO-TAILED SWALLOWTAIL has been known to inhabit
the Yosemite region since 1930, when B. A. Thaxter and Edna Banta,
Yosemite Field School members, wrote: "Among the swallowtails found
here we took Papilio daunus, the largest western species, a bright
yellow butterfly with black markings and two tails on each hind
wing.” Their brief report, "Some Butterflies and Moths of the Yose-
mite Valley Region,” is on file at the Yosemite Museum, but their
two specimens are without accompanying data. For information on
exact locality and date we are indebted to Oakley Shields, who twice
encountered it in August at Jerseydale, where it came to thistles.
Life Zones: Upper Sonoran, Transition. Plant Communities: All
at moderate elevations. Host Plants: SALICACEAE: Populus (Cot-
tonwood), Salix (Willow). LAURACEAE: Umbellularia (Laurel).
OLEACEAE: Fraxinus (Ash). ROSACEAE: Amelanchier (Service
Berry), Prunus (= Cerasus) (Choke-Cherry). RUTACEAE: Ptelea
(Hop Tree).
Ilia. Papilio eurymedon eurymedon Luc.
b. Papilio eurymedon albanus F. & F.
THE PALE SWALLOWTAIL is more frequently encountered on
dry hillsides than is the nearly related and equally abundant Papilio
rutulus. While the food plant adheres strictly to the lower zones, the
strong wings of the butterfly carry it to the tops of the highest peaks.
A form or possible subspecies occurring at the higher altitudes, smaller
in size and darker in color, is known as P. e. albanus F. & F.
Life Zones: Upper Sonoran, Transition, straying higher. Plant
Communities: Chaparral, Foothill Woodland, and the chaparral-like
under story of most Coniferous Forests. Host Plants: RHAMNACEAE:
Rhamnus (= Frangula) calif ornicus (Coffee Berry), R. crocea (Red
Berry), Ceanothus (Snow Bush).
2(i):i-^6, ip6}
YOSEMITE BUTTERFLIES
63
112. Parnassius clodius haldur Edw. PI. II, fig. e
THE BALDUR PARNASSIAN flies to the very rim of the preci-
pitous walls of Yosemite Valley, but does not descend into the valley
itself. In the Research Reserve it was particularly abundant, flying
over thickets of Castanea (Chinquapin) and Holodiscus (Ocean Spray),
and pausing occasionally to sip nectar from yellow Sene do, a composite.
While the fir belt generally defines its habitat, it occasionally strays
higher among Lodgepole Pine and Mountain Hemlock. A few indi-
viduals are without the black-margined secondaries and suggest the
parent species, P. dodius Men.
Life Zones: (upper Transition), Canadian, lower Hudsonian
(Arctic- Alpine ) . Plant Communities: Red Fir Forest; Lodgepole Pine
Forest; exceptionally, Alpine Fell- Fields. Host Plants: CRASSUL-
ACEAE: Sedum (Stonecrop) .• ERICACEAE: Vacdnium ( Bilberry
SAXIFRAGACEAE: Saxifraga (Saxifrage).
113. Parnassius phoebus hehrii Ed^v. PL III, fig. f
BEHR’S PARNASSIAN is well established on the eastern slope of
the Sierra Nevada, as befits its affinities with the Great Basin fauna. The
arid eastern exposures of many ridges west of the Sierran divide, with
the persistent association of Juniper and Sedum (Stonecrop), have
served to lure this butterfly westward through Tioga Pass to Yosemite
Creek: It also flies in the Potentilla fruticosa zone of the Alpine Rock
Garden sub-association of the Alpine Fell-Fields. The larvae and pupae
of P. s. behrii were first described from specimens secured by the
senior author and Edmund Godwin at Rock Creek Lake.
Life Zones: Canadian, Hudsonian, Arctic- Alpine. Plant Commun-
ities: Juniper Woodland, Alpine Fell-Fields. Host Plants CRASSUL-
ACEAE: Rhodiola rosea (Western Roseroot), Sedum ohtusatum, S.
stenopetalum (Stonecrop). SAXIFRAGACEAE: Saxifraga (Saxi-
frage) .
HESPERIIDAE
114. Epargyreus darus (Cram.)
Syn. Epargyreus tityrus ( Fabr. )
THE SILVER-SPOTTED SKIPPER is a widely distributed species
recognizable by its large size and the conspicuous silvery spot on the
under side of each hind wing. It frequents the flowers of California
Buckeye, Aesculus calif ornica, a well known tree of the western foot-
hills, and is favored by plantings of Locust as an ornamental.
Life Zones: Upper Sonoran. Plant Communities: Foothill Wood-
land, Yellow Pine Forest. Host Plants: LEGUMINOSAE: Rohinia (In-'
troduced Black Locust), and other legumes.
115. Thorybes pylades (Scud.)
THE NORTHERN DUSKY- WING is the low-elevation species
of this genus in California. It is larger but has smaller spots on the
forewing than the following Thorybes mexicana nevada, and the male
has a costal fold.
64
GARTH AND TILDEN
/. Res. Lepid.
Life Zones: Upper Sonoran. Plant Communities: Chaparral, Foot-
hill Woodland, (Yellow Pine Forest). Host Plants: LEGUMINOSAE:
Trifolium (Clover), and other legumes.
116. Thorybes mexicana nevadaScwd.
THE NEVADA DUSKY-WING is the alpine and subalpine
Thorybes of the Sierra Nevada, flying to 11,000 feet. Smaller and
with larger spots than the preceding T, pylades, the male has no costal
fold and the spots are outlined with a dark rim.
Life Zones: Canadian, Hudsonian, Arctic- Alpine. Plant Commun-
ities: Alpine Fell-Fields, Subalpine Forest (Glacial Moraine). Host
Plants: Unknown. LEGUMINOSAE: Amorpha calif ornica (False In-
digo) for the parent species, according to Comstock and Dammers
(1933).
117. Thorybes diversus Bell PL I, fig. a
BELL’S DUSKY-WING is easily confused with The Northern
Dusky-Wing, Thorybes pylades, being of similar size, but lacks the
costal fold in the male. A denizen of grassy openings in the forest belt,
it shuns the broad, open meadows.
Life Zones: Transition. Plant Communities: Openings in several
types of Forest. Host Plants: Not known.
118. Pyrgus ruralis (Bdv.)
THE TWO-BANDED SKIPPER prefers openings in the forest
rather than damp meadows, and is best distinguished by its darker
color, even checkering, and montane habitat. Flying close to the
ground, it is inconspicuous and easily overlooked.
Life Zones: Canadian. Plant Communities: Openings in Red Fir
and Lodgepole Pine Forest. Host Plants: MALVACEAE: Sidalcea
( Checkerbloom, Wild Hollyhock). ROSACEAE: Potentilla (Cinque-
foil) suggested.
119. Pyrgus communis (Grote)
THE COMMON CHECKERED SKIPPER is abundant at low
elevations, often near habitations where introduced mallow grows. It
may be recognized by the conspicuous flight habit, bluish-gray and
white checkered pattern, and proximity to human dwellings.
Life Zones: Upper Sonoran, Transition. Plant Communities: Sev-
eral. Favors weedy growth around cultivated areas. Host Plants:
MALVACEAE: especially Malva (Mallow) and Sidalcea (Checker-
bloom). The larvae draw the leaves together with silken threads.
120. Heliopetes ericetorum (Bdv.)
THE LARGE WHITE SKIPPER is a rapid flier, difficult to cap-
ture, and hence a rarity. The female is more heavily marked than the
male and resembles the foregoing Pyrgus communis.
Life Zones: Upper Sonoran (lower Transition). Plant Commun-
ities: Chaparral, Foothill Woodland, "Mountain Misery” {Chamae-
2(i):i-96, 1963
YOSEMITE BUTTERFLIES
65
batia folios a) belt of cut-over stands of Yellow Pine Forest. Host
Plants: AMARANTHACEAE: Amaranthus (Pigweed). MALVA-
CEAE: Mdvastrum (Apricot Mallow).
121. Erynnis persius ( Scud . )
THE PERSIUS DUSKYWING is a small, dark insect with few
contrasting markings. Long misidentified in the west, it has been
shown by Burns (unpublished doctoral dissertation) to be common
to California at moderate elevations.
Life Zones: (Upper Sonoran) Transition, Canadian. Plant Com-
munities: (Foothill Woodland), Yellow Pine and Fir Forests, both
White and Red. Host Plants: Best regarded as unknown. Early rec-
ords are untrustworthy.
122. Erynnis pacuvius lilius ( Dyar )
DYAR’S DUSKYWING, as identified by John Burns, is common
at intermediate elevations. Gin Flat, Crane Flat, and similar locations
are its usual haunts. One of us (Tilden) found it common on Cow-
chilla Mountain, outside the park, in June, 1954.
Life Zones: (upper Transition) Canadian. Plant Communities:
Chaparral -like understory of montane forests. Host Plants: Unknown.
RHAMNACEAE: Ceanothus (Snow Bush, Wild Lilac) for Colorado
pacuvius, as reported by Donald Eff (personal communication).
123. Erynnis propertius (Scud. & Burg.)
THE "PROPERTIUS DUSKWING is the largest and commonest
of our Duskywings. The front wings have few spots and present a
slightly "grizzled” appearance, rather than a solid black effect. It
frequents edges and openings in the forest at moderate elevations.
Life Zones: (Upper Sonoran) Transition, Canadian. Plant Com-
munities: Chaparral, Foothill Woodland, Yellow Pine and other Coni-
ferous Forests. Host Plants: FAGACEAE: Quercus (Oak), especially
Live Oaks of various species.
124. Erynnis zarucco funeralis (Scud. & Burg.)
THE FUNEREAL DUSKYWING is the most easily recognized of
our Duskywings. The front wing is pointed and the fringes of the
hind wing are white. There is an indistinct brownish patch on the
outer third of the forewing that is lacking in the following Erynnis
tristis; the males also have a tibial tuft.
Life Zones: (Lower Sonoran) Upper Sonoran. Plant Communi-
ities: Valley Grasslands, Foothill Woodland (Chaparral). Host Plants:
LEGUMINOSAE: Hosackia ( = Lotus) scoparia, Medicago (Alfalfa),
and other legumes.
125. Erynnis tristis ( Bdv. )
THE SAD DUSKYWING may be expected to occur in the oak
woodlands and foothills, although it has not yet been found in the
Yosemite region. Slightly smaller than the preceding Erynnis z. fun-
66
GARTH AND TILDEN
/. Res. Lepid.
eralis, and with the forewing not noticeably pointed, it lacks the
brown patch on the forewing and the tibial tuft.
Life Zones: Upper Sonoran. Plant Communities: Foothill Wood-
land, Oak Woodland. Host Plants. FAG ACE AE: Quercm (Oak).
126a. Hesperia harpalus harpalus (Edw.) PL II, fig. g
b. Hesperia harpalus yosemite Leuss.
THE HARPALUS SKIPPER is the characteristic Hesperia of the
eastern declivity of the Sierra Nevada and the adjacent Great Basin
area surrounding Mono Lake. It is rather pale above, and shows well
developed silver spots below, especially in the females. THE YOSE-
MITE SKIPPER is the subspecies that occurs on the western slope
of the Sierra. It is smaller and more brightly colored than normal
harpalus and the spotting of the under sides of the hind wings tends
to be small or to disappear. At high elevations there occurs a "blend-
zone” form that is intermediate. The literature treats the Great Basin
form as H. idaho ( Edw. ) , but there seems no reason for retaining this
name.
Life Zones: Transition, Canadian. Of the "blend-zone” population:
Hudsonian ( Arctic- Alpine ) . Plant Communities: Not noticeably se-
lective. Host Plants: POACEAE: (Grasses).
127. Hesperia miriamae MacNeill PL III, fig. m
MIRIAM’S SKIPPER is the most alpine of our skippers, being
found entirely above tree line on Mono Pass and on the summits of
the higher peaks, such as Unicorn. It is a small, dark species, with
large, irregular spots on the under surface of the hind wings. But
recently described (MacNeill, 1959), it is still a rarity, and will continue
to be so due to the inaccessibility of its habitat.
Life Zones: Arctic- Alpine. Plant Communities: Alpine Fell-Fields.
Host Plants: POACEAE (Grasses) , presumptively. Actually, unknown.
128. Hesperia nevada (Scud.)
THE NEVADA SKIPPER is a Great Basin species found on the
eastern Sierra slope from Mono Lake to elevated areas above Tioga
Pass. A small species with irregular macular bands on the under side
of the hind wings, it frequents open rocky areas at high elevations.
Life Zones: Arid Transition, Canadian, Hudsonian. Plant Com-
munities: Juniper Woodland, Subalpine Forest, Alpine Fell-Fields.
Host Plants: POACEAE: Grasses.
129. Hesperia juba (Scud.)
THE YUBA SKIPPER is not limited by elevation, being found
sparingly across the Sierra crest from the western foothills to the Great
Basin. A large species of powerful flight, it may be recognized by
the fuscous border of the front wing, which is sharply limited and does
not grade into the rufous ground color. The under side of the wings
is greenish brown to dark brown and the spots are large and usually
white.
s(i):i--96> 1963
YOSEMITE BUTTERFLIES
67
Life Zones: Unlimited. Plant Communities: Apparently not re-
stricted. Ubiquitous. Host Plants: POACEAE: (Grasses), probably.
130. Ochlodes sylvanoides (Bdy.)
THE WOODLAND SKIPPER is one of the commonest skippers,
ranging across the mountains from the western foothills to the Great
Basin and eastward to Colorado. It is a moderate-sized, orange-brown
species, with the stigma of the male touching the dusky border of the
wing. An unwary species, it may be found fairly swarming on roadside
flowers. The Mono Lake population is slightly paler, but differs from
Colorado napd Edw., a subspecies with which it has been confused.
Life Zones: Upper Sonoran, Transition (Canadian). Plant Com-
munities: Several. Not restricted within its life zones. Host Plants:
POACEAE : ( Grasses ) .
131. Ochlodes agricola ( Bdv. )
THE FARMER flies from May in the western foothills to July at
higher elevations, but is absent from the east side of the Sierra Nevada.
It frequents streamsides, openings in lush woodlands, and overgrown
roadsides. Smaller and darker than the preceding Ochlodes sylvanoides,
it shows pale translucent spots in the forewing, at the end of the stigma
in the male, and near the end of the cell in the female. It is seldom
found in wet meadows or dry fields.
Life Zones: Upper Sonoran, Transition. Plant Communities: Open-
ings in Forests. Riparian and Roadside Associations. Host Plants:
POACEAE: (Grasses).
132a. Polites sabuleti sabuleti (Bdv.)
b. Polites sabuleti tecumseh (Grin.) PL III, fig. d
THE SANDHILL SKIPPER is found in the Yosemite area, in one
form or another, entirely across the Sierran crest, from the western
foothills to the Great Basin. About the size of The Woodland Skipper,
Polites s. sabuleti has the under surface of the hind wings tan, the spots
yellowish and extending onto the wing veins, which are also yellow.
At higher elevations it is replaced by THE TECUMSEH SKIPPER,
which fairly swarms in the subalpine meadows adjacent to Tioga Pass.
It is much smaller, darker, brighter, and more greenish than normal
Polites sabuleti. Its flight season, July to September, is long for an
alpine species, perhaps indicating two broods.
Life Zones: Upper Sonoran, Transition for P. sabuleti; Canadian,
Hudsonian, Arctic- Alpine for P. s. tecumseh. Plant Communities: Not
restrictive for P. sabuleti; Subalpine Meadow, Alpine Meadow and
Grassland for P. s. tecumseh. Host Plants: CYPERACEAE: Car ex
filifolia (Sedge). Also recorded as LEGUMINOSAE: Trifolium
monanthum (Clover). The latter, if correct, is a departure from the
food preferences of this group, which are characterized as feeding on
Monocotyledons.
68
GARTH AND TILDEN
/. Res. Lepid.
133. Polites sonora {Scud.)
THE SONORAN SKIPPER is the usual mid-summer skipper of
intermediate elevations in Yosemite National Park. Resembling the
Woodland Skipper, Ochlodes sylv amides, in size and appearance, it
is a darker, more olivaceous rufous, with a thicker, more complex
stigma. Below, the hind wings are olivaceous tan with a light basal
spot and a band of light spots across the wing. A somewhat lighter
phase occurs in the higher portions of the Great Basin, as at Mono
Lake.
Life Zones: Arid Transition, Canadian, Hudsonian. Plant Com-
munities: Meadowlands, Freshwater Marsh, Alkali Sink; openings in
forests of several kinds. Host Plants: Unknown.
134.. Amhlyscirtes vialis (Edw.)
THE ROADSIDE SKIPPER is a widely distributed species, found
in the cooler parts of the United States and southern Canada from
California to Maine. In the Yosemite region it is a rarity. A dull dun-
colored species, of small size, with a few small specks near the tip of
the forewing, purplish gray shades on the under surface, and checkered
fringes, it frequents cool, moist spots, such as meadow edges and for-
est clearings, at elevations of from 4,000 to 7,000 feet.
Life Zones: Transition, Canadian. Plant Communities: Not known
for the Yosemite region. Host Plants: POACEAE: (Grasses) in the
eastern United States; has not been reared in the west.
LITERATURE CITED
BADE, WILLIAM F. 1924. The life and letters of John Muir. In two volumes.
Vol. I: vii 399 pp. Houghton Mifflin, Boston & New York.
BAUER, DAVID L. 1961. Tribe Melitaeini, Checker Spots, Crescents, and
Patch Butterflies, pp. 127-146. In: Ehrlich, Paul R., and Anne H. How
to know the Butterflies. 262 pp. Wm. C. Brown, Dubuque, Iowa.
BROWN, F. MARTIN. 1954. Colorado Butterflies. Part I. Satyridae.
Proc. Denver Mus. Nat. Hist., No. 3: 1-32.
1955. Studies of Nearctic Coenonympha tullia ( Rhopalocera, Satyri-
dae.) Coenanympha tullia inornata Edw. Bull. Amer. Mus. Nat. Hist.,
105 (4): 363-409.
CLENCH, HARRY. 1961. Tribe Theclini, Hairstreaks, pp. 177-220. Tribe
Lycaenini, The Coppers, pp. 220-228. /«:Ehrlich, Paul R., and Anne H.
How to know the Butterflies. 262 pp. Wm. C. Brown, Dubuque, Iowa.
COMSTOCK, JOHN A. 1927. Butterflies of California. 334 pp., 63 pis.
Author, Los Angeles.*
DAVENPORT, DEMOREST. 1941. The butterflies of the satyrid genus
Coenonympha. Bull. Mus. Comp. ZooL, Harvard, 87(4): 215-349,
pis. 1-10.
DICE, LEE R. 1943. The biotic provinces of North America, vii + 78 pp.,
1 map Univ. Michigan Press, Ann Arbor.
DOS PASSOS, C. F., and L. P. GREY. 1947. Systematic catalogue of
Speyeria ( Lepidoptera, Nymphalidae) with designations of types and
fixations of type localities. Amer. Mus. Novitates, No. 1370: 1-30.
DOWNEY, JOHN C. 1961. Tribe Plebejini, The Blues, pp. 228-242. In:
Ehrlich and Ehrlich. How to Know the Butterflies. 262 pp. Wm. C.
Brown, Dubuque, Iowa.
196}
YOSEMITE BUTTERFLIES
69
EMMEL, THOMAS C., and JOHN F. EMMEL. 1962. Ecological studies of
Rhopalocera in a High Sierran community — Donner Pass, California.
1. Butterfuly associations and distributional factors. /. Lepid. Soc., 16:
23-44.
— 1963. Larval food-plant records for six western Papilios. /. Res.
Lepid., 1(3): 191-193.
ERIKSEN, C. H. 1962. Further evidence of the distribution of some boreal
Lepidoptera in the Sierra Nevada. J. Res. Lepid., 1(1): 89-93. ^
EHRLICH, PAUL R., and ANNE H. EHRLICH. 1961. How to know the
butterflies, vi^- 262 pp., 524 figs. Wm. C. Brown, Dubuque, Iowa.
GARTH, JOHN S. 1935a. Butterflies of the Boundary Hill Research Reserve,
Yosetmite National Park, California. Bull. Sou. Calif. Acad. Sci., 33:
131-135.
— — “ 1935b. Butterflies of Yosemite National Park. Ibid., 34: 37-75.
GRINNELL, JOSEPH, and HARVEY M. HALL. 1919. Life-zone indi-
cators in California. Proc. Calif. Acad. Sci., 9(2) : 37-67.
— , and TRACY T. STORER. 1924. Animal life in the Yosemite.
xviii +752 pp., 62 pis., 65 figs. Univ. Calif. Press, Berkeley.
GUNDER, JEAN D. 1929. The genus Euphydryas Scud, of Boreal America
(Lepidoptera, Nymphalidae) . Pan-Pac. Ent., 6(1) : 1-8, pis. 1-12, maps 2.
HALL, HARVEY M., and CARLOTTA C. HALL. 1912. A Yosemite flora.
vii + 282 pp. Paul Elder, San Francisco.
HEMMING, FRANCIS. 1934. Generic names of Holarctic butterflies. 184 pp.
British Museum (Natural History), London.
HIGGINS, L. G, 1953. Butterfly collecting in U.S.A. Entomologist, 86(9) :
207-210,
HOVANITZ, WILLIAM 1940. Ecological color variation in a butterfly and
the problem of "protective coloration.” EcoL, 21 (3): 371-380.
— — 1943. Hybridization and seasonal segregation in two races of a butter-
fly occurring together in two localities. Biol. Bull. 85: 44-51.
— — - — “. 1950. The biology of Colias butterflies. I. The distribution of the
North American species. Wasmann J. Biol., 8(1): 49-75.
- — - — 1963. Evidence for the species distinction of Pieris protodice and
Pieris occidentalis. J. Res. Lepid. 2: in press.
KLOTS, ALEXANDER B. 1936. The interrelationships of the species of
the genus Lycaena Fabricius (Lepidoptera, Lycaenidae). Bull. Brooklyn
Ent. Soc., 31(4): 154-170.
— 1951. A field guide to the butterflies of North America, east of the
Great Plains, xvi -f 349 pp., 40 pis., 9 figs. Houghton Mifflin, Boston.
LINDSEY, ARTHUR W. 1942. A preliminary revision of Hesperia. Den-
ison Unit’. Bull., Jour. Sci. Labs., 37 : 1-50, pis. 1-6.
— , E. L. BELL, and R. C. WILLIAMS. 1931. Hesperioidea of North
America. Ibid., 26: 1-142.
MATTHES, FRANCOIS E. 1930. Geologic History of the Yosemite Valley.
Geol. Surv. Professional Paper I6O: 1-137, 52 pis.
MacNEILL, C. DON. 1959. A new species of Hesperia from California
(Lepidoptera: Hesperiidae) . Wasmann J. Biol., 17(1): 89-94.
McDUNNOUGH, J. 1938. Check list of the Lepidoptera of Canada and the
United States of America. Part 1. Macrolepidoptera. Mem. Sou. Calif.
Acad. Sci., 1: 1-275-
MUNZ, PHILIP A., and DAVID D. KECK. 1949. California plant com-
munities. El Aliso, 2(1): 87-105. 1950. (Same title) — -Supplement.
Ibid., 2(3) ; 199-202.
NABOKOV, VLADIMIR. 1949. The Nearctic members of the genus
Lycaeides Hbl. Bull. Mus. Comp. ZooL, Harvard, 101: 479-541.
OPLER, PAUL, and JERRY A. POWELL. 1961. Taxonomic and distribu-
tional studies on the western components of the Apodemia mormo complex
(Riodinidae) . J. Lepid. Soc., 15(3): 145-171.
70
GARTH AND TILDEN
/. Res. Lepid.
TILDEN, J. W.. 1959. The butterfly associations of Tioga Pass. Wasmann J.
Biol., 17(2): 249-271.
VESTAL, A. G. 1914. Internal relations of terrestrial associations. Amer.
Nat., 48: 413-445.
WILLIAMS, C. B. 1930, The migration of butterflies. Oliver and Boyd,
Edinburgh.
ZIEGLER, J. BENJAMIN. I960. Preliminary contribution to a redefinition
of the genera of North American hairstreaks (Lycaenidae) north of
Mexico. /. Lepid. Soc., 14: 19-23.
YOSEMITE BUTTERFLY RECORDS
The documented records on which the textual material rests have been
segregated from the body of the article (Appendix I) because of its length;
however these localities are of great importance to specialists interested in the
specific locations where these species may be found.
West-slope records precede east-slope records and are separated from
them by a period. Each locality is followed by all subsequent records from
that locality, first by the same observer, then by other observers. Localities
are separated by a semi-colon. Apart from these intended departures an
exact chronology is maintained.
Following the abbreviated locality and date (day, MONTH, and year),
the initials of the responsible observer are given. These are John S. Garth
(JSG), Edmund D. Godwin (EDG), Allan Oakley Shields (AOS), J. Wilson
Tilden (JWT), and Keith Trexler (KT). Others who contributed an occas-
ional record are named with the appropriate citation.
LOCALITIES AND ABBREVIATIONS
To conserve space in the body of the list a standard set of abbreviations
has been adopted, (Appendix II) following the practice established by
entomological reports of a similar nature (Garth, 1935b) : Localities repre-
sented extend from Briceburg and Buck Meadows at the western approaches
to Yosemite National Park to Mono Lake at the eastern Gateway, and from
Conway Summit on the north to Deadmans Summit on the south along U. S.
Highway 395. (A few localities, such as Cathay and Oakhurst on the west
side and Bridgeport and Casa Diablo Hot Springs on the east side, transcend
even these limits.) Some are in the National Park, some are in the surrounding
National Forests, while some are outside of either. Localities outside Yosemite
National Park are identified by National Forest (NF), of if outside the
National Forest, by California County ( Co. ) . Each locality is characterized
by Life Zone and Plant Community; the division into west slope and east
slope suffices to indicate the Biotic Province. Elevations (7214, 10,500)
are in feet and are taken from U. S. D. I. Geological Survey Topographical
Maps of Yosemite (1948) or of adjacent Quadrangles. These are accurate
to the nearest 100-foot contour except in the case of mountain peaks, lake
levels, or places of habitation, where precise figures (BM) can be given.
2(i);i-96, 196}
YOSEMITE BUTTERFLIES
71
APPENDIX I
SOME PLANTS ON WHICH YOSEMITE BUTTERFLIES FEED AS LARVAE
o = oviposition observed; 1 - larva taken from; r = reared to adult
on plant named; ? - doubtful record; * = feeding on more than one
plant
AMARANTHAGEAE
Amaranthus
ASCLEPIADACEAE
Asclepias
Pigweed
Milkweed
*Heliopetes ericetorum
Danaus plexippus o, 1, r
Danaus strigosus
BETULACEAE
Alnus
BORAGINACEAE
Amsinckia
Cryptantha
Cynoglossum
Mertensia
Alder
*Papilio rutulus
*Polygonia t. rusticus
Fiddle-Neck *Vanessa cardui
Nievitas *Vanessa cardui
Hound's Tongue *S trymon melinus
Mertensia Euphydryas c. sierra
1. r
1* r
1, r
CAPPARIDACEAE
Cleome
Isomeris
CAPRIFOLIACEAE
Symphoricarpus
CARYOPHYLLACEAE
S tellaria
CHENOPODIACEAE
Bee Plant *Pieris protodice
Bladder Pod *Pieris beckeri
Chickweed
*Nathalis iole
o, 1, r
o, 1, r
Snow Berry * Euphydryas chalcedona o, 1, r
Atrlplex
Chenop odium
COMPOS ITAE
Salt Bush *Brephidium exilis
Lamb's Quarters *Brephidium exilis
Anaphalis
Antennaria
Artemesia
Aster
Carduus
Everlasting
Everlasting
Mugwort
Aster
Thistle
^Vanessa virginiensis 1
*Vanessa virginiensis 1
*Vanessa virginiensis
*Melitaea acastus
?Melltaea d. malcolmi
Phyciodes campestris
*Phyciodes mylitta o, 1, r
* Vanessa cardui o, 1, r
72
GARTH AND TILDEN
/. Res. Lepid.
Cirsium
Dyssodia
Gnaphalium
Helenium
Silybum
CONVOLVULACEAE
Cuscuta
CORNACEAE
Cornus
CRASSULACEAE
Rhodiola
Sedum
CRUCIFERAE
Arabia
Barbarea
Brassica
Caulanthus
Dentaria
Erysimum
Lepidium
Radicula
Raphanus
Sisymbrium
Stanleya
Streptanthus
Thlaspi
Thistle
(none)
Cud Weed,
Everlasting
Sneeze Weed
Milk Thistle
*Phyciodes mylitta
^Vanessa cardui
*Nathalis iole
^Vanessa virginiensis
*Nathalis iole
*Phyciodes mylitta
o, 1, r
o, 1, r
1
Dodder
*?Callophrys iroides
Dogwood
^Celastrina argiolus 1, r
Roseroot *Parnassius 2* behrii 1
Stonecrop *Parnassius c. baldur
*P amass jus 2* behrii 1
Rock Cress
Winter Cress
Mustard
Rock Cabbage
Milk Maids
Wall Flower
Pepper Grass _
Water Cress
Radish
Hedge Mustard
Desert Plume,
Prince's Plume
Jewel Flower
Anthocharis lanceolata
*Anthocharis sara
^•Euchloe ausonides
^Euchloe creusa
*Pieris occidentalis
*Pieris protodice
*P_ieris sisymbrii
*Pieris napi
* Anthocharis sara
*Pieris beckeri
*Pieris napi
*Pieris protodice
*Pieris rapae
*Pieris sisymbrii
*Pieris napi
^•Euchloe ausonides
^•Pieris occidentalis
*Pieris protodice
^Pieris rapae
*Pieris napi
*Pieris protodice
* Anthocharis sara
*Euchloe ausonides
*Pieris beckeri
*^Pieris occidentalis
*Pieris protodice
*Pieris sisymbrii
*Pleris beckeri
*Euchloe creusa
*Pieris sisymbrii
*Pieris protodice
o
o, 1, r
o, 1, r
o
O, 1, r
o, 1, r
o, 1, r
o, 1, r
o, 1
o, 1
o
o
o.
1
o
1.
r
Penny Cress
2(i):i-96, 196}
YOSEMITE BUTTERFLIES
73
ERICACEAE
Arbutus
Arctostaphylos
Rhododendron
Vaccinlum
FAGACEAE
Quercus
GERANIACEAE
Erodlum
GUTTIFERAE
Hypericum
LABIATAE
Hyp t Is
LAURACEAE
Umbellularia
LEGUMINOSAE
Amorpha
Astragalus
Dalea
Glycyrrhiza
Hosackia
( - Lotus)
Madrono
Manzanita
Azalea
Bilberry,
Huckleberry*
*Callophrys iroides
*Callophrys iroides
*Celastrina argiolus
*Polygonia _f . rusticus
*Polygonia zephyrus
*Callophrys iroides
Colias behrii
*?Pamassius c. baldur
1,
r
o,
1,
Oak
Limenitis b. californica
*Celastrina argiolus
Erynnis propertius
Erynnis tristis
Habrodais grunus
?Strymon auretorum
*?Strymon californica
Filaree
*Nathalis iole
St. John's Wort *Strymon melinus
Bee Sage
* Strymon melinus
Laurel *?Papilio multicaudatus
False Indigo Colias eurydice
Thorybes mexicana
Loco Weed, *Colias eury theme
Rattle Weed
*Everes amyntula
*Everes comyntas
*Glaucopsyche lygdamus
*Hemiargu8 isolus
*Lycaeides melissa
*Plebe jus acmon
*Strymon behrii
*Strymon melinus
(Includes Smoke *Leptotes marina
Tree)
Liquorice *Lycaeides melissa
*?Celastrina argiolus
*Colias eury theme
*?Erynnis z. funeral is
*Glaucopsyche lygdamus
*Hemiargus isolus
*Lycaeides melissa
*Plebe jus acmon
o, 1
o, 1
o
o, 1, r
o
74
GARTH AND TILDEN
/. Res. Lepid.
*Plebeius saepiolus o
*Strymon behrii
*Stirymon melinus
(2nd brood)
o, 1, r
Lathyrus
Sweet Pea
*Everes amyntula
*Glaucopsyche lygdamus
Lupinus
Lupine
*?Celastrina argiolus
*Glaucopsyche lygdamus
?Lycaeides a. anna
*Lycaeides melissa
Phaedrotes piasus
Plebeius icarioides
Strymon fuliginosum
*Strymon melinus
o, 1
o, 1 , r
Medicago
Alfalfa
*Colias eury theme
*Erynnis z. funeralis
*Leptotes marina
*Lycaeides melissa
0, 1, r
Melilotus
Sweet Clover
*Colias eury theme
o
Rob ini a
Black Locust
Epargyreus clarus
Therraopsis
False Lupine
*Colias eurytheme
Trifolium
Clover
Colias philodice hagen
*Colias eurytheme
*Everes comyntas
^Hemiargus isolus
*Plebeius saepiolus o
?Polites sabuleti
Thorybes pylades
li o, 1, r
o
(1st brood)
Vicia
Vetch
*Colias eurytheme
*Everes amyntula
*Everes comyntas
*Glaucopsyche lygdamus
o, 1
o
LORANTHACEAE
Arceuthobium
Dwarf Mistle-
toe
Callophrys iohnsoni
Callophrys spinetorum
1, r
Phoradendron
Mistletoe
Atlides halesus
1, r
MALVACEAE
Lavatera
Tree Mallow
*Vanessa carye
Malva
Mallow
*Strymon melinus
*Vanessa carye
*PyrKus communis
r
o, 1, r
1, r
Malvastrum
Apricot Mallow
*Heliopetes ericetorum
Sida
Alkali Mallow
*Vanessa carye
Sidalcea
Checkerbloom, *Pyrsus coranunis
Wild Hollyhock
*?Pyrgus ruralis
*Vanessa carye
o, 1, r
Sphaeralcea
Bush Mallow, *Vanessa carye
Apricot Mallow
ONAGRACEAE
Gayophytum
*Lycaena helloides
2(i):i-^6, 1^6}
YOSEMITE BUTTERFLIES
75
OLEACEAE
Fraxlnus
Pit; ACE AE
Libocedrus
Pinus
Pseudotsuga
PLANTAGINACEAE
Plantago
POLYGONACEAE
Erlogonum
Oxytheca
Polygonum
Rumex
RHAMNACEAE
Ceanothus
Rhamnus
(= Frangula)
ROSACEAE
Amelanchier
Cercocarpus
Ash
*Papllio multicaudatus
Incense Cedar
Pine
Douglas Fir
Callophrys nelsoni, probably
Callophrys eryphon
^Neophasia menapia
*Neophasia menapia
Plantain
Euphydryas edltha o, 1, r
^Precis o. evarete o, 1, r
Buckwheat
(none)
Knot Weed
Dock, Sorrel
Apodemia mormo
Callophrys dumentorum o
?Callophrys lemberti
Lycaena gorgon o
Lycaena heteronea
? Lycaena nivalis
Philotes battoides
Philotes enoptes o
*Plebeius acmon
?Plebeius lupini
*Lycaena helloides
Philotes speciosa
*Lycaena helloides
^Lycaena mariposa
*S trymon melinus
*Lycaena editha o
^Lycaena helloides o
^Lycaena mariposa
? Lycaena hypophlaeas
? Lycaena rubidus
Lycaena xanthoides o
Snow Bush
Cascara, Buck-
thorn, Coffee
Berry
^Callophrys iroides
*Celastrina argiolus
Erynnis pgcuvis
*Papilio eurymedon
Nymphalis califomica
S trymon califomica
S trymon saepium
*Papillo eurymedon
1, r
1, r
1
1, r
1, r
1, r
Service Berry *Papillo mul t icaudatus
Mountain Maho- S trymon adenos,tomatis 1, r
gany
S trymon califomica 1, r
S trymon saepium
76
GARTH AND TILDEN
/. Res. Lepid.
Potentilla
(= Horkelia)
Cinquefoil
*Lycaena editha
*?Pyrgus rural is
Prunus
(= Cerasus)
Choke Cherry
*Limenitis lorquini
*Papilio multicaudatus
*Papilio rutulus
Purshia
Antelope Bush
Strymon fuliginosum
Spiraea
Spirea
*Celastrina argiolus
RUTACEAE
Ptelea
SALICACEAE
Hop Tree
*Papilio multicaudatus
1
1, r
Populus
Salix
Poplar
Willow
*Limenitis lorquini 1
*L imenitis w. nevadae
*Nyinphalls antiopa o, 1, r
*Papilio multicaudatus
*Papilio rutulus o, 1, r
*Limenltls lorquini
*Limenitis w. nevadae
*Nymphalis antiopa o, 1, r
*Papilio multicaudatus
*Papilio rutulus
*?PolyRonia _f. rusticus
Strymon dry ope 1, r
S trymon sylvinus 1, r
SAP INDACE AE
Aesculus Horse Chestnut *Celastrina argiolus 1, r
(includes Buckeye)
SAXIFRAGACEAE
Glossularia Gooseberry
Ribes Currant
SaxigraRa Saxifrage
SCROPHULARIACEAE
*Lycaena arota o, 1, r
*PolyRonia _f . rusticus
*PolyRonia zephyrus
*Lycaena arota
*Pamassius c. baldur
*Pamassius 2* behrii
Castilleja
Cordylanthus
Diplacus
Mimulus
Penstemon
Scrophularia
Paint Brush
Bird's Beak
Sticky Monkey
Flower
Monkey Flower
Penstemon
Figwort, Bee
Plant
*Melitaea acastus
*Melitaea leanira
*Melitaea leanira
*Euphydryas chalcedona o, 1, r
*Euphydryas chalcedona
^Precis o. evarete
*Euphydryas chalcedona
*Euphydryas chalcedona o, 1, r
2(i):i-^6, 1963
YOSEMITE BUTTERFLIES
77
ULMACEAE
Ulmus
UMBELLIFERAE
Daucus
Foenlculum
Heracleum
Pteryxia
URTICACEAE
Boehmeria
Parietaria
Urtica
VIOLACEAE
Viola
Elm
^Nymphalis antiopa o, 1, r
Wild Carrot
Fennel, Anise
Cow Parsnip
*Papilio zelicaon
^Papilio zelicaon
^'Papilio zelicaon
Papilio indra
o,
1, r
1, r
1, r
False Nettle
Pellitory
Nettle
^'Vanessa atalanta
^Vanessa atalanta
Nymptialis milberti
Polygonia satyrus
^Vanessa atalanta
1, r
1, r
Violet
Argynnis (all)
Boloria epithore
THREE ANOMALOUS SITUATIONS
While the regularity of zonal progression and the exclusiveness of biotic provinces have
been emphasized, certain irregularities occur that are of sufficient interest to the lepidopterist
to warrant considration. The first of these may be found at Tioga Pass, where the loosely
consolidated moraines provide a situation duplicating the alpine rock garden. Within a few
feet of these moraines may be seen flying the Arctic-Alpine butterflies, Oeneis chryxus ivallda
and Euphydryas editha nubigena, species that normally occur above timber line, a thousand
or more feet higher. The explanation is believed to be found in drainage, both of air that
funnels through the pass and cools the ground sufficiently to permit the growth of Alpine
Willow, and of water that enables these slightly elevated rock piles to become snow-free with
the first rays of summer sun, and to flower a month before the Alpine Fell-Fields above.
A second paradoxical situation occurs at timberline, as along the Gaylor Lakes trail
above Tioga Pass, where at an approximate elevation of 10,500 feet may be found Artemesian
species that normally occur at Mono Lake, 4,000 feet below. Cercyonis oeta, Lycaena rubidus,
L. heteronea, and Hemiargus isolus are found in a sage-brush environment much like Mono
Lake, but with Artemesia rotbrockii , rather than A. tridentata, as the sage, and with Eriogonum
latifolium and Aplopappus sp. as the plant associates. The butterflies are not "fly-ups” from
below, but appear to be permanently established on these brushy summits. Here the tie to |
plant community, the Sagebrush Scrub, seems stronger than the restriction to a particular life
zone or altitudinal belt. Thus one may, after observing "Arctic-Alpine” butterflies at Tioga
Pass, nominally Hudsonian, ascend a thousand feet and observe "Upper Sonoran” butterflies
flying at tree-line!
While normally the Sierran Crest divides the Californian biotic province from the
Artemesian, a series of lesser crests to the westward support Western Juniper on their east-
ern slopes, and with it the Juniper-Woodland butterflies, Parnassius phoebus behrii, Euchloe
ausonides coloradensis, and Hesperia nevada. It is therefore not necessary to travel to the
eastern slope of the Sierra Nevada to observe Great Basin butterflies; many of these have
infiltrated the mountain passes and may be found in favored situations as far west as Yosemite
Creek, well within the central portion of the park.
78
GARTH AND TILDEN
/. Res. Lepid.
APPENDIX II
YOSEMITE RECORDS
SATYRIDAE
la, Coenonympha tullia californica West. & Hew. El Portal 28- VI- 33
(JSG); Mather 12- VII- 56 (JSG), 14-V-61 (JWT); Crane Flat 18- VIII- 57
(JWT); Jerseydale 30-V-59 and 11- VI-61 (JWT), 26- VI-59 (JSG), 30-
VIII-60 (AOS); Briceburg 21-III-61 (KT); Cliff House 9-VI-61 (JWT).
b. Coenonympha tullia mono Burdick Mono Lake 3 to 5- VII- 38 and 23-
VI-62 (JWT), 30-VI-'5Frnd 12- VI- 62 (AOS); W of Lee Vining 26 to 28-
VI- 61 (AOS, common).
2. Cercyonis pegala ariane (Bdv. ) Mono Lake 17-VII-57 (JWT), 16 to 20-
VII- 58 and 13- VII- 60 (AOS); 10 miles E of Mono Lake 24- VII- 59 (AOS).
3. Cercyonis silvestris (Edw. ) El Portal 28- VI- 33 (JSG), 10- VI- 61
(JWT); Pate 23-VII-34 (EDG); Benson 24-VII-34 (EDG); Hetch-Hetchy
13 to 15-VII-56 (JSG); Indian Flat 30-V-59 (JWT, seen), lO-VI-61
(JWT); Jerseydale 26- VI-59 (JSG), ll-VI-61 (JWT); Cathay ll-VI-61
(JWT).
4. Cercyonis oeta( Bdv. ) Tioga Pass 19-VIII-52 (JWT); Crest W of Tioga
Pass 6-VIII-57, 31- VII to 20- VIII- 58 and 23 to 28- VII- 60 (AOS); Upper
Gaylor Lakes l-VIII-58 (JSG), 20-VIII-58 (AOS); Pilot Peak 31-VIII-
58 (JWT). Bodie 18- VII-57 (AOS); June Lake 14 to 22-VIII-57 (JSG);
Mono Lake 6-VII-58 (AOS), 23- VI-62 (JWT); Warren Creek 2 1 to 28-
VII-58, ll-VIII-60 and 26-VI-61 (AOS); Agnew Pass and Thousand Is-
land Lake 5-VIII-59 (AOS).
5. Oeneis chryxus ivallda (Mead) Dana, W slope 7- VII- 3 1 (JWT); Flor-
ence 5-VIII-33 (JSG); Lyell 6-VUI-33 (JSG); Dana 8- VIII- 33 (JSG);
Slide l-VIII-34 (EDG); Pilot Peak 16-VIII-52 and 18- VII-57 (JWT);
Crest W of Tioga Pass 12 to 15-VII-57 and 9-VII to 19-VIII-58 (AOS);
Tioga Pass lO-VII-58 (AOS); Helen Lake 18 to 29-VII-58 (AOS); Cocks-
comb 26-VII-58 (AOS); Dana, N slope 3 1-VII-58 (AOS); Upper Gaylor
Lake l-VIII-58 (JSG); Vogelsang Pass 3-Vm-58 (JSG); Bert Lake
9- VIII- 58 (AOS); Unicorn 12- VIII- 58 (AOS).
DANA IDA E
6. Danaus (Danaus) plexippus (Linn.) Camp 9, Yosemite Valley, 23-VII-
33 (JSG); Yosemite Creek 7-VII-56 (JSG); Mather 11 and 12- VII- 56
(JSG); Crane Flat 19- VII- 57 (JWT); Darrah, Jerseydale, and Indian
Flat 29 and 30-V-59 (JWT, seen); Jerseydale and Cathay 11- VI-61
(JWT). Mono Lake 30- VI-41 (JSG, seen), 23-VI-62 (JWT).
7. Danaus (Tasitia) gilippus strigosus (Bates) Mono Lake 30-VI-30 (JWT,
seen); l6-Vn-58 (AOSK
NYMPHALIDAE
8. Argynnis (Semnopsyche) cybele leto Behr Camp 9, Yosemite Valley,
23- VII to 2 -VIII- 3 3 (JSG)'
9. Argynnis (Speyeria) nokomis apacheana Skin. Mono Lake 13- VIII- 50
and 15- VIII- 51 (JWT), 22 to 26- VIII- 57, 25-VIII-58, and4-VIII-60
(AOS); Gull Lake 14 to 28-VIII-57 (JSG), 2-IX-58 (JWT).
10 a. Argynnis (Speyeria) zerene zerene Bdv. Wawona 20- VII- 33 (JSG);
Jerseydale 21 to 24- VI-56, 19- VI-58, and 9 to 13- VI- 62 (AOS); Mather
11 to 15-VII-56 (JSG); Hetch-Hetchy Summit 13-VII-56 (JSG); Aspen
2(i):i~^6, 1963
YOSEMITE BUTTERFLIES
79
Valley 14-VII-56 (JSG); Yosemite Creek Trail 17-VII-56 (JSG); Crane
Flat I9-VIII-57 (JWT).
b. Argynnis (Speyeria) zerene malcolmi Comst. Agnew Pass 13- VIII- 57
and 5- VIII- 59 (AOS);“M^^I^Lake 26-VIII-57 (AOS); Lundy 20- VII- 58
(AOS); Warren Creek 28-Vni-58 (AOS); Casa Diablo 2-IX-58 (JWT).
11a. Argynnis (Speyeria) callippe inornata Edw. Jerseydale 2 1- VI- 56, 23-
VI-57, 5- VII- 60, Tl to 23 -VI- 61 and^ to 13- VI- 62 (AOS); El Portal
30-V-58 (JWT); Oakhurst, Madera County, 24. VI- 62’ (JWT).
b. Argynnis (Speyeria) callippe nevadensis Edw. Mono Lake 15- VII- 38,
30 -VI- 50 and 23-VI-62 (JWT), 7-VII-58 (AOS); Above Crowley Lake
29- VI-41 (JSG); Bodie 18- VII- 57 (AOS); 5 miles N of Mono Lake 28- VI-
61 (AOS).
12. Argynnis (Speyeria) egleis Bdv. Eagle Peak l-VII-33 (JSG); Reserve
l6^Vn-33 (TsG); Dana 8-Vni-33 (JSG); Pate 23-VII-34 (EDG); Yose-
mite Creek 9- VII- 56 (JSG); Mather 12- VII- 56 (JSG); Hetch-Hetchy Sum-
mit 13-VII-56 (JSG); Smoky Jack 4-Vn-57 (JWT); Gin Flat 12 to 19-VII-
57 (JWT); Lembert Dome 24- VII- 57 (AOS); Crest W of Tioga Pass 6-
Vni-57, 19-VIII-58, DVni-59 and 14-VIIto 2-VIII-60 (AOS); Crane Flat
18- VIII-57 (JWT); Pilot Peak 30 to 31-VIII-58 (JWT); Unicorn 27- VII-
59 (AOS); Glen Aulin 25-VII-60 (AOS). June Lake 14-Vni-57 (JSG);
Lundy 20-VII-58 (AOS); Warren Creek 21- VII- 58 (AOS).
13. Argynnis (Speyeria) atlantis irene Bdv. Coldwater 3- VIII- 34 (EDG);
Tenaya 4- Vni-34 (EDG); Aspen 14- VII- 56 (JSG); 5 miles W of Lower
Ottoway Lake 19- VIII- 61 (AOS).
14. Argynnis (Speyeria) hydaspe Bdv. Camp 19, Yosemite Valley 5-Vn-
33 (JSG); Ledge 9-VII-33 (JSG); Glen Aulin lO-VIII-33 (JSG); Pate 23-
Vn-34 (EDG); Buck Meadow 4- VII- 54 (JWT); Crane Flat 4- VII- 54 (JWT);
Jerseydale 28- VI-56, 15 to 16- VI- 58, 18- VI and 4- VH- 60 (AOS), 26- VI-
59 (JSG); Mather 12-Vn-56 (JSG); Hetch-Hetchy Summit 13 to 17-Vn-
56 (JSG).
15. Argynnis (Speyeria) mormonia arge Stkr. Tioga Pass 13-VIII-29 (JSG),
I4-VII-31, 12 to 13-Vin-50, and“T6^ VIII- 57 (JWT), 4-Vni-57, 19-VIII-
58 and l-VIII-59 (AOS); Dana 8-Vin-33 (JSG); Coldwater 3-Vm-34
(EDG); Crane Flat 19-VII to 19-VIII-57 (JWT); Tuolumne Meadows 18-
Vn to 14-Vni-57 and 3-IX-58 (JWT), 15 to 21-VIII-58 (AOS); Pilot
Peak 15- VIII- 57 and 3- VIII- 58 (JWT); Dana, W slope l6-Vni-57 and
30- VIII-58 (JWT); Lyell Fork lO-VIII-58 and 3-VIII-59 (AOS); Gaylor
Lakes 31-VIIIto 3-IX-58 (JWT); Elizabeth Lake 27-VII-59 (AOS).
Warren Creek ll-Vni-60 (AOS).
16. Boloria (Clossiana) epithore (Edw.) Tioga Pass 7-VII-31 (JWT); Eagle
Pi"ak I-VIlTFTjSG); LedgT 9-Vn-33 (JSG); Reserve 16-VII-33 and 7
to lO-VII-56 (JSG); Yosemite Creek 7 to 17-VII-56 (JSG); White Wolf
10- VII-56 and 24- VI-59 (JSG); Aspen Valley 14-Vn-56 (JSG); Crane
Flat l9-Vn-57 and lO-VI-61 (JWT), 24- VI-61 (AOS, abundant); Tenaya
11- VII-58 (AOS); Badger Pass 23- VI-59 (JSG); Glacier Point 23-VI-59t
(JSG); Tamarack 24- VI- 59 (JSG).
17a. Euphydryas chalcedona chalcedona (Dbldy. & Hew, ) El Portal 28- VI-
33 (JSG), 14-V-$1 (JWT) ; Glacier 9-VII-33 (JSG); Jerseydale 16 to
19- VI-57, 15 to 19-VI.58 and 9 to 13-VI-62 (AOS); Darrah and Jersey-
dale 30-V-59 (JWT); Indian Flat 30-V-59 and 14-V-61 (JWT).
b. Euphydryas chalcedona sierra (Wgt. ) Return Creek ll-VII-31 (JWT);
Eagle Peak l-VII-33 (JSG); Reserve 14-VII-33 (JSG).
80
GARTH AND TILDEN
/. Res. Lepid.
18a. Euphydryas editha rubicunda Hy. Edw. El Portal 8-V~32 (R. G. Wind),
14-V-61 (JWT); Jerseydale 15 to 18- VI-58 and 9 to 13- VI- 62 (AOS);in-
dian Flat 30-V-59, 27-V-60 and 13-V-61 (JWT).
b. Euphydryas editha nubigena (Behr) Tioga Pass 7-VII-31, 19-VII-52
and 18-Vn-57 (JWT), 29-VII-56, 8 to 13-VII-57, 9- VII to 24- VIII- 58,
20 to 23-VII-60 and 25-VI-61 (AOS), l-Vni-58 (JSG); Pilot Peak 18-VII
to 15-Vm-57 (JWT); Helen Lake 18 to 29-VII-58 (AOS); Gaylor Lakes
Trail 19-VII-58 (JWT); Dana, W slope 20-Vn-58 (JWT); Rafferty Creek
3-VIII-58 (JSG); Vogelsang Lake 3-VIII-58 (JSG).
c. Euphydryas editha monoensis Gund. Mono Lake, near Lee Vining 4-
VII-28 and 30-VI-50 (JWT); 19-VI-54 (JSG), 30- VI to 16-VII-58, 13-Vn-
60 and 26 to 27-VI-61 (AOS).
19. Melitaea (Chlosyne) damoetas malcolmi Comst. Dana, W slope 17-
VII-3 1 (JWT); Lyell 6- VIII- 33 (JSG); Dana 8- VIII- 3 3 (JSG); Pilot Peak
16- VIII-52 and 18-Vn-57 (JWT); Upper Gaylor Lake 3 1-VII-57 and 9
to 19-VII-58 (AOS), l-VIII-58 (JSG); Crest W of Tioga Pass 9-Vnto
3 1-VII-58, 14-VIIto 2-VIII-60 and 25-VI-61 (AOS); Helen Lake 18-VII-58
(AOS); Dana, N slope 3 1- VII to 24-Vin-58 and 6 to 8- VIII- 60 (AOS);
Bert Lake 9- VIII- 58 (AOS). East above Saddlebag Lake 28- VII -58
(AOS).
20. Melitaea (Chlosyne) acastus Edw. Mono Lake 5-VII-38 and 30-VI-50
(JWT), 30-VI-41 (JSG), 30-VI-58 (AOS).
2 1a. Melitaea (Chlosyne) palla palla Bdv. Eagle Peak 1- VII- 33 (JSGU Ledge,
Glacier 9-VII-33 (JSG); Reserve l6-Vn-33 (JSG); Yosemite Creek 7 to
17- VII-56 (JSG); White Wolf lO-VII-56 (JSG); Indian Flat 30-V-59, 27-
V- 60 and 13-V-61 (JWT); El Portal 14-V-61 (JWT); Jerseydale 21-VI-
61 (AOS).
22. Melitaea (Chlosyne) hoffmanni Behr Reserve 14 to 18-VII-33 (JSG);
Pate 23-VII-34 (EDG); Benson 24-VII-34 (EDG); White Wolf 10-VII-
56 (JSG).
23. Melitaea (Chlosyne) leaniraF. & F. Ledge 9-VII-33 (JSG); Tamarack
Flat 3- VIII- 54 (JWT); Darrah 30-V-57 (JWT); Indian Flat 30-V-59
and 13-Vto lO-VI-61 (JWT); El Portal 14-V-61 (JWT); Jerseydale
22-VI-61 (AOS).
24a. Phyciodes (Phyciodes) campestris campestris (Behr) Museum 5-Vn-
33 (JSG); Mather 12- VII- 56 (JSG), 10- VI-61 and 30-VI-62 (JWT);
Hetch-Hetchy 13-VII-56 (JSG); Jerseydale 23-VI-61 (AOS).
b. Phyciodes (Phyciodes) campestris montana (Behr) Reserve l6-Vn-33
(JSG); Dana 8-Vni-33 (JSG); Coldwater 3- VIII- 34 (EDG); Tamarack
Flat 3-VII-54 (JWT); Aspen Valley 14-Vn-56 (JSG); below Helen Lake
19-VII-60 (AOS). Thousand Island Lake 4-VIII-59 (AOS).
c. Phyciodes (Phyciodes) campestris, intermediate form Mono Lake 30-
VI- 41 and 19-VI-54 (JSG), 17- VII- 57 and 23-VI-62 (JWT), 30-VI to 20-
VII- 58, 13-VII-60, 26 to 27- VI-61 and 12-VI-62 (AOS); Huntoon Public
Camp, Bridgeport 19- VI- 54 (JSG); June Lake 14 to 29- VIII- 57 (JSG);
Warren Creek 21 and 28- VII- 58 (AOS); W of Lee Vining 26 to 27- VI- 61
(AOS).
25. Phyciodes (Phyciodes) mylitta (Edw. ) Museum 5-VII-33 (JSG); Benson
24-VII-34 (EDG); Yosemite Creek 9-VII-56 (JSG); Aspen Valley 14- VII-
56 (JSG); Crane Flat 18-Vm-57 (JWT), 24- VI-59 (JSG); Tioga Pass 1-
IX-57 (AOS); Tenaya ll-Vn-58 (AOS); Glacier Point 23-VI-59 (JSG);
Jerseydale 26-VI-59 (JSG), 30- VIII- 60 (AOS); El Portal 2 l-IH- 6 1 (KT),
2(i):i.s6, 1963
YOSEMITE BUTTERFLIES
81
14-V-61 (JWT);Yosemite Valley 19”IV~6l (KT); Indian Flat 13-V-61
(JWT); Mather 10- VI-61 (JWT); Warren Creek ll-VIII-60 (AOS).
26. Polygonia satyrus (Edw. ) Museum 5-VII-33 (JSG). Mono Lake 20-VI
t^25-VIII-58T^27-VI-6l (AOS).
27. Polygonia faunus rusticus (Edw.) "Yosemite" VI-26 (E, O. Essig).
Specimen in Yosemite Museum in 1933.
28. Polygonia zephyrus (Edw.) Reserve 14-Vn-33 (JSG); Yosemite
^eek 9-'viLT6 (JSG); Aspen Valley 14-Vn-56 (JSG); Jerseydale 17-
VII-56 and 16-VI-58 (AOS); Dana, W slope 20-VII-58 (JWT); Tioga
Pass 20-VIII-58 (AOS); Upper Gaylor Lake 20-VIII-58 (AOS); Dana, N
slope 31-VII to 24-VIII-58 and 8-Vin-60 (AOS); Badger Pass 23-VI-59
(JSG); Tuolumne Grove 24- VI-59 (JSG). Mono Lake 30-. VI-41 (JSG).
29. Nymphalis californica (Bdv. ) Yosemite Valley 24- VI to 5-VII-33
(JSG); Ledge 9-VII-33 (JSG); Darrah, Jerseydale, and Indian Flat 30-
V- 59 (JWT); Bridal Veil Creek 23- VI-59 (JSG); White Wolf 24- VI-59
(JSG); Cliff House 9- VI- 61 (JWT); Crane Flat, Gin Flat, and Mather
lO-VI-61 (JWT) (outbreak numbers); El Portal lO-VI-61 (JWT); Jer-
seydale and Cathay 11- VI-61 (JWT). Donohue Pass 4-VIII-59 (AOS).
30. Nymphalis milberti furcil-lata (Say) Lyell 6- VIII- 33 (JSG); Tioga
Pass 8-VIIt^-Vin-57, 1- VIII- 58, and 25- VI- 61 (AOS); Crest W of
Tioga Pass 1 to 19-VIII-58 (AOS); Lyell Base Camp 9-VIII-58 (AOS);
Dana, N slope 24-VIII-58 (AOS). Mono Lake 30-VI-41 (JSG), 26-VI-
61 (AOS); Warren Creek 29- VIII- 57 (AOS); Agnew Meadows 9- VIII-
58 (AOS).
31. Nymphalis antiopa (Linn.) Research Reserve 8-'VdI-56 (JSG, seen).
Mono Lake 30- VI-41 (JSG, seen).
32. Vanessa atalanta (Linn.) "Yosemite" VI-26 (E. O. Essig).
33. Vanessa virginiensis (Dru. ) Research Reserve 14 to 18-VII-33 and
T0^^Yn^6 (JSG); Yosemite Creek Trail 9-VII-56 (JSG); Mather 10-
VI- 61 (JWT).
34. Vanessa cardui (Linn.) Research Reserve 14 to 18-VII-33 (JSG);
YosT mit e“cT^ Trail 9- VII- 56 (JSG).
35. Vanessa carye (Hbn. ) Research Reserve 14 to 18-VII-33 (JSG);
Hatch- Hetchy 12- VII- 56 (JSG). Huntoon Public Camp, Bridgeport 19-
VI- 54 (JSG).
Precis orithya evarete (Cram.) Ledge Trail 9-VII-33 (JSG); Research
Reserve 14 to 19-VII-33 and 8 to lO-VII-56 (JSG); Yosemite Creek
Trail 9“ VII- 56 (JSG); Mather Hand 12-VII-56 (JSG), lO-VI-61 (JWT);
Jerseydale 17~VI-60 (AOS); El Portal 4-IV- 61 (KT); Indian Flat 13- V-
61 (JWT); Cathay 11- VI- 61 (JWT). Mono Lake 13- VII- 60 (AOS).
37. Limenitis (Limenitis) weidemeyerii nevadae (B. Er B.) Mono Lake 30-
VI-41 (jSG)7“30-VI-567jWTT7 30-VI to 20TviI-58 and 26- VI- 61 (AOS).
Limenitis (L. ) weidemeyerii nevadae X Limenitis (L.) lorqmni (Hy-
brid) Mono“Lak7l7^VIII- 5 1 (JWT), 19- VI- 54 (JSgJ, 7 to 20- VII- 58,
25-VII-59 and 27-VI-61 (AOS).
38. Limenitis (Limenitis) lorquini Bdv. Yosemite Valley VII- 33 (JSG);
Pate ValIiy'23-VII-34 "(EDCFAspen Valley 14- VII-56 (JSG); Jersey-
dale 16-VI-58 (AOS); Darrah and Jerseydale 30-V-59 (JWT, seen);
Indian Flat 30-V-59 (seen) and 13-V-61 (JWT); Cliff House 9-VI-61
(JWT); Mather lO-VI-61 (JWT). Mono Lake 30-VI-58 (AOS).
39. Limenitis (Adelpha) bredowii californica (Butl.) Museum 5-Vn-33
(JSG); Camp 9, Meadow 23-VII-33 (JSG); Mather 12 to 15- VII- 56 (JSG).
82
GARTH AND TILDEN
/. Res. Lepid.
10- VI-61 (JWT); Hetch-Hetchy 13 to 15-VII-56 (JSG); Aspen Valley 14-
VII-.56 (JSG); Crane Flat 24- VI-59 (JSG); Jerseydale 26-VI-59 (JSG),
11- VI-61 (JWT), 9 to 13-VI-62 (AOS); Darrah, Jerseydale, and Indian
Flat 30-V-59 (JWT); El Portal 14-V-61 (JWT); Cliff House 9-VI-61
(JWT).
RIODINIDAE
40a. Apodemia mormo mormo F. & F. Little Yosemite 3-VIII-33 (JSG).
Mono Lake 22 to 26- VIII- 57 and 20- VII- 58 (AOS); June Lake 22-Vm-
57 (JSG); Conway Pass, N of Mono Lake 21-VIII-60 (JWT).
b. Apodemia mormo tuolumnensis Opler & Powell Pate Valley 23-VII-34
(EDG, as A. m. virgulti).
LYCAENIDAE
41. Habrodais grunus (Bdv. ) Columbia Point 14-VII-33 (JSG); Pohono
Trail 23- VII- 33 (JSG); Benson 24- VII- 34 (EDG); Hetch-Hetchy 13- VII-
56 (JSG); Jerseydale 21 to 23-VI-61 (AOS).
42. Atlides halesus estesi Clench Museum 5-Vni-34 (collector unknown);
Yosemite Creek Trail 9- VII- 56 (JSG).
43. Strymon (Strymon) melinus pudica (Hy. Edw. ) Museum 8-VII-33
(JSG); Mather 12- VII- 56 (JSG); Darrah 29-V-59 (JWT).
44. Strymon (Satyrium) fuliginosum semiluna (Klots) Mono Lake 30- VI- 50
(JWT), 3 O-VI-58 ^ A'osyi
45. Strymon (Satyrium) behrii (Edw.) Mono Lake 30-VI-50 and 23-VI-62
(JWT), 30-VI to 20- VII-58 and 27- VI- 61 (AOS); W of Lee Vining 18-
Vn-57 and 24- VI-58 (AOS); Bodie 18-VII-57 (AOS); Mono Craters
23- VII- 57 (AOS); Warren Creek 26- VI-61 (AOS). 5 miles N of Mono
Lake 28- VI- 61 (AOS).
46. Strympn (Satyrium) auretorum (Bdv.) El Portal 28- VI-33 (JSG), 10-
VI- 61 (JWT); Briceburg and Mariposa 26- VI- 54 (JWT); Hetch-Hetchy
Road 13 to 15-VII-56 (JSG); Jerseydale 21 and 22- VI-58 (AOS); Darrah
29- V-59 (JWT); Cathay ll-VI-61 (JWT).
47. Strymon (Satyritim) saepium (Bdv.) El Portal 28- VI-33 (JSG); Pate
23-VII-34 (EDG); Hetch-Hetchy Road 13-VII-56 (JSG); Gin Flat 17-Vni-
57 (JWT); Jerseydale 21 and 22- VI- 58 (AOS); Darrah and Indian Flat
30- V-59 (JWT); Cathay 28-V-60 and ll-VI-61 (JWT); Cliff House 9-
VI- 61 (JWT). Warren Creek 28- VII and 28- VIII- 58 (AOS); Gull Lake
2-IX-6I (JWT).
48. Strymon (Satyrium) adenostomatis (Hy. Edw.) Mariposa 26-VI-54
(JWT); Jerseydale 21 and 22-VI-58 (AOS); Darrah 29-V-59 (JWT).
49. Strymon (Satyrium) sylvinus (Bdv.) Museum 8-Vn-33 (JSG); Bear
Creek LodJiT^^I- 5T(TWT); Hetch-Hetchy Road 13-VII-56 (JSG);
Jerseydale 2 1- VI- 6 1 <(AOS). Mono Lake 5-VII-38 (JWT).
Strymon (Satyrium) californica (Edw.) Old Village 12-VI-32 (JWT);
Museum 5-VII-33 (JSG); Pate 23- VII- 34 (EDG); Big Oak Flat Road
2-VII-54 (JWT); Mirror Lake 2- VII- 54 (JWT); Tamarack Flat 4-Vn-
54 (JWT); Hetch-Hetchy Road 13 to 15- VII- 56 (JSG); Mather 15- VH-
56 (JSG); Jerseydale 15 to 22-VI-58, 17- VI- 60 and 2 1 to 23- VI- 6 1
(AOS); Darrah 30-V-59 (JWT); El Portal 14- V and 10- VI- 6 1 (JWT);
Cathay ll-VI-61 (JWT). Mono Lake 5- VII- 38, 30-VI-50, 17-Vn-57,
and23-VI-62 (JWT), 30- VI-58 and 27- VI-6 1 (AOS); Lee Vining 24-
VI-58 (AOS); Warren Creek 2 1- VII-58 (AOS); 5 miles N of Mono
2(i):i-s6, t$6}
YOSEMITE BUTTERFLIES
83
Lake 28- VI- 61 (A”OS).
Strymon (Satyrium) dryope (Edw. ) Mono Lake 20- VII- 58 and 4- VIII- 60
(AOS); Bridgeport 15- VIII- 52 (JWT).
52. Callophrys (Callophrys) dumetorum (Bdv. ) Briceburg 21-III- 6 1 (KT);
El Portal l6TlvT6l"~(JWTTr“ Mather 14-V-61 (JWT).
53. Callophrys (Callophrys) lemberti Tilden Reserve 16-VII-33 (JSG);
Pil^ Peak I'o'-VII and l6^Wl-5’2 (JWT); White Wolf lO-VII-56 (JSG);
Crest W of Tioga Pass 9-VII-58 and 25-VI-61 (AOS), 20-VII-58
(JWT); Tioga Pass lO-VII-58 (AOS). Warren Creek 12-VI-62 (AOS).
54. Callophrys (Mitoura) spinetorum (Hew.) Tamarack Flat 4- VII- 54
(JWT); JersT^^daTI” I?- VI- 57Ti7d 27-VIIto 3- VIII- 62 (AOS); Tioga
Road 24- VI- 59 (JSG).
55. Callophrys (Mitoura) johnsoni (Skin.) Jerseydale 27 and 28-VII-56,
^VIII- 56 anT^T^Vn-I^lAOS) .
56. Callophrys (Mitoura) nelsoni (Bdv.) Musetim 5-VII-33 (JSG); Reserve
Ti toT8-'^L3T7jSG)rijetch-Hetchy 13-VII-56 (JSG); Jerseydale 30-
V- 59, 27-V-60, and ll-VI-61 (JWT), 26- VI-59 (JSG), 9 to 13-VI-
62 (AOS); Crane Flat 24- VI-59 (JSG); Highway 120 and Hetch-Hetchy
Road 14- V- 61 (JWT); Cliff House 9-VI-61 (JWT); Mather 10- VI- 61
(JWT).
57. Callophrys (Incisalia) doudoroffi windi (Clench) Indian Flat 16-IV-61
__ _
58. Callophrys (Incisalia) iroides (Bdv.) Ledge 9-VII-33 (JSG); Crane
Flat 3- VII- 54 (JWT); “jerseydale 19- VI-58 and 21 to 23- VI-61 (AOS),
29-V-59 (JWT); Darrah 29-V-59 (JWT); Glacier Point 23-VI-59
'(JSG); El Portal 4-IV-61 (KT); Mather lO-VI-61 (JWT).
59. Callophrys(Incisalia)eryphon (Bdv.) Reserve 9 to 14-VII-33 (JSG);
TamaraclTFlat 4- VII- 54 (JWT); Smoky Jack 4- VII- 54 (JWT);
Mather 12-VII-56 (JSG), lO-VI-61 (JWT); Jerseydale 16-VI-58
(AOS), 29-V-59 (JWT); Tenaya Canyon 11- VII- 58 (AOS). Mono
Lake 27- VI- 61 (AOS).
60a. Lycaena (Tharsalea) arota arota (Bdv.) Museum 8-VII-33 (JSG);
^te 23-VJl34~l¥DG)r^e77i7dale 29-VI to 27-VII-56, l-VIII-58,
24-Vin-60, 23-VI-61 and 28-Vn-62 (AOS),
b. Lycaena (Tharsalea) arota virginiensis (Edw.) Mono Lake 17-VII-57
TjWtTT” 2 6‘^^IL 5 7T rt^O-VII-58 and 13- VII- 60 (AOS).
61. Lycaena (Lycaena) gorgon (Bdv.) Darrah 29-V-59 (JWT); Indian
^at 30-vT59~^ ll-V to lO-VI-61 (JWT); El Portal 14-V-61 (JWT).
62. Lycaena (Lycaena) heteronea Bdv. Lyell Base Camp 26- VII- 57 (AOS);
Tioga Pas”i~4rvni-“5T"lAOS); Merced Lake Trail 4- VIII- 58 (AOS);
Gaylor Lakes Trail 3-IX- 58 (JWT); Dana, N slope 8-Vni-60 (AOS).
Warren Creek 7- VIII-57 and 2 1- VII to 28-Vm-58 (AOS); Lundy 20-
VII- 58 (AOS); Mono Lake 13- VII- 60 (AOS). Thousand Island Lake
4-Vm-59 (AOS),
63. Lycaena (Lycaena) xanthoides (Bdv.) Mather 11 and 12-VII-56 (JSG),
(JWT);” HSt^^Hitchy 13- VII- 56 (JSG); Aspen Valley 14-
VII- 56 (JSG).
64. Lycaena (Lycaena) rubidus (Behr) Mono Lake 27-VII-37 (JWT), 30-
VI- 41 (JSG), 30- VI to 7^11-58 and 27- VI-61 (AOS); Gaylor Lakes
Trail 16-VIII-52 (JWT); Warren Creek 7 to 29-Vin-57, 21-VIIto
28-VIII-58 and ll-FIII-60 (AOS); Agnew Pass 5-Vm-59 (AOS); W
of Lee Vining 2 8- VI- 61 (AOS).
84
GARTH AND TILDEN
/. Res. Lepid.
65. Lycaena (Lycaena) editha (Mead) Kuna 7- VIII- 33 (JSG); Dana 8- VIII-
33 (JSG): Kerrick 27- VII- 34 (EDG); Snow Lake 29- VII- 34 (EDG);
Tamarack Flat 4- VII- 54 (JWT); Tioga Pas s 16 to 17- VII- 57 (JWT),
14- VII to 24- VIII- 58 aTid 14- VIII- 60 (AOS); Gin Flat and Crane Flat
19-VII-57 (JWT); Rafferty Creek 21-VII-57 (AOS); Tioga Meadow
15- VIII-57 (JWT); Pilot Peak 15- VIII- 57 and 3 1 - VIII- 58 (JWT);
Dana, W slope 17- VIII- 57 and 20- VIII- 58 (JWT); Tuolumne Meadows
15 to 21- VIII-58 (AOS), 3-IX-58 (JWT); Gaylor Lakes Trail 30-VIII
to 3-IX-58 (JWT); Lyell Base Camp 3-VIII-59 (AOS); below Agnew
Pass 5-VIII-59 (AOS). Mono Lake 30- VI-41 (JSG); Bodie 18-VII-57
(AOS); Warren Creek 21-VII-58 (AOS).
66. Lycaena (Lycaena) mariposa Reak. Kuna 7-VIII-33 (JSG); Slide 29-
VII- 34 (EDG); Dana, W slope 13-VIII-50, 15-VIII-51, 16-VIII-57
and 30- VIII-58 (JWT); Lyell Base Camp Trail 26-VII-57 and lO-VHI-
58 (AOS); Tioga Pass 4-VIII-57, 20- VIII- 58 and 1- VIII- 59 (AOS);
Pilot Peak 15-VIII-57 (JWT); Elizabeth Lake 27- VII to 22- VIII-58
and 27- VII- 59 (AOS); Merced Lake Trail 4- VIII- 58 (AOS). Agnew
Meadow 13- VIII- 57 (AOS).
67. Lycaena (Lycaena) nivalis (Bdv. ) Tioga Pass 14-VII-31 (JWT), 20-
VIII- 58 (AOS); Glacier 9- VII- 33 (JSG); Reserve 14-VII-33 (JSG);
Kuna 7- VIII- 33 (JSG); Benson 24-Vn-34 (EDG); Kerrick 27- VII-
34 (EDG); Gaylor Lakes Trail 15-VIII-51, 6- VIII- 53 and 19- VII to 3-
IX- 58 (JWT); Smoky Jack 3- VII- 54 (JWT); Yosemite Creek Trail
9- VII- 56 (JSG); Dog Lake Trail 19- VII- 56 (JSG); Rafferty Creek
Trail 21- VII- 57 and 24- VII- 58 (AOS); 3-VIII-58 (JSG); Tenaya Can-
yon ll-VII-58 (AOS); Crest W of Tioga Pass 4 to 18- VIII- 57, 20-
VIII- 58 and 20 to 23- VII- 60 (AOS); Lyell Fork Meadows 10- VIII- 58
and 3- VIII- 59 (AOS); Dana, N slope 24- VIII- 58 (AOS). Warren
Creek 21-VII-58 (AOS).
68. Lycaena (Lycaena) helloides (Bdv.) El Portal 28- VI-33 (JSG); Mather
12- VII- 56 (JSG), Mono Lake 5-VII-38 (JWT).
69. Lycaena (Lycaena) phlaeas hypophlaeas (Bdv.) Lyell 6-Vin-33 (JSG);
Dana, N slope 6 to 13- VIII- 60 (AOS).
70. Lycaena (Lycaena) cupreus (Edw. ) Tioga Pass 7-VII-31, 13-Vni-50,
15- VIII- 51, 16- VII to 16- VIII- 52, 6-VIII-53, and 18- VII to 16-VIII-57
(JWT), 7 to 15-Vn-57, 9-VII to 24- VIII-58, 15-VII-60 and 25-VI-61
(AOS); Kuna 7-VIII-33 (JSG); Dana 8-Vin-33 (JSG); Snow Lake 29-
VII-34 (EDG); Dog Lake Trail 19-VII-56 (JSG); Pilot Peak 18- VU
to 15-VIII-57 (JWT); Cockscomb Peak 26- VI-58 (AOS); Tuolumne
Meadows 4-VII-58 (AOS); Crest W of Tioga Pass 9-VII to 1- VIII- 58
(AOS); Helen Lake 18 to 29- VII- 58 (AOS); Elizabeth Lake 27-VII-
58 (AOS); Upper Gaylor Lake 1- VIII- 58 (JSG); Rafferty Creek Trail
3- VIII- 58 (AOS).
71. Leptotes marina (Reak.) Mono Lake 30- VI-41 (JSG), 30- VI-50
(JWT, seen) and 23-VI-62 (JWT).
72. Brephidium exilis (Bdv.) E above Saddlebag Lake 28-VII-58 (AOS);
Mono Lake 23-VI-62 (JWT).
73. Hemiargus (Echinargus) isolus (Reak.) Tioga Pass 9-Vnto 7-Vin-58
(AOS); Crest W of Tioga Pass 9- VII- 58 (AOS); Upper Gaylor Lake
14-Vn-58 (AOS); Rafferty Creek Trail 3- VIII- 58 (JSG, AOS); Vo-
gelsang Lake 3-VIII-58 (JSG, AOS); Gaylor Lakes Trail 31-VIIIto
3-IX-58 (JWT); Tuolumne Meadows 3-IX-58 (JWT). Mono Lake 20-
VU-58 (AOS); Lee Vining 24-VII-58 (AOS); Warren Creek 28-VII-
2(i):i-96, 1963
YOSEMITE BUTTERFLIES
85
58 (AOS).
74. Lycaeides argyrognomon anna (Edw. ) Conness Creek ll-VII-31
■(JWT); Dana 8- VIII- 33 (JSG);" Kerrick 27- VIE 34 (EDG); Coldwater
3- VIII- 34 (EDG); Tenaya 4- VIII- 34 (EDG); Crane Flat 2- VII- 54
and 19-Vn-57 (JWT); Hetch-Hetchy Summit 13-VII-56 (JSG); Aspen
Valley 14-VII-56 (JSG); Gin Flat 19-VII-57 (JWT); Tuolumne Mea-
dows 14-VIII-57 (JWT); Badger Pass 23-VI-59 (JSG).
75. Lycaeides melissa inyoensis Nab. Reserve 16-VII-33 (JSG); Ben-
son 24-Vli3riEbG); Helen Lake 18- VII- 58 (AOS); Cockscomb
Peak 26-VII-58 (AOS); Upper Lyell Base Camp lO-VIII-58 and 3-
VIII- 59 (AOS). Mono Lake 20- VII- 58 (AOS), 23- VI- 62 (JWT); E
above Saddlebag Lake 28-VIL58 (AOS); Warren Creek ll-Vni-60
and 26- VI-61 (AOS); Conway Sximmit 23-VI-62 (JWT).
76. Plebejus (Agriades) glandon podarce (F. & F. ) Tioga Pass 7-VII-31,
TTand 14 ->111-50, 15 - VIII- 517 16 - VIH- 52, 6 and 7- VIII- 53, and 16-
VIII-57 (JWT), 1 to 24-Vm-58 and 23- VII-60 (AOS); Eagle Peak 1-
Vn-33 (JSG); Reserve 18-VII-33 and lO-VII-56 (JSG); Kuna 7-VIII-
33 (JSG); Kerrick 27-VII-34 (EDG); Tuolumne Meadows 16-VII-57
(JWT); Dana, W slope 17-VIII-57 and30-Vni-58 (JWT); Elizabeth
Lake 27-VII-58 (AOS); Helen Lake 29-VII-58 (AOS); Upper Gaylor
Lake l-VIII-58 (JSG); Bert Lake 9-VIII-58 (AOS); Unicorn Peak
23-Vni-58 (AOS); Bridal Veil 23- VI- 59 (JSG); White Wolf 24- VI-
59 (JSG); Dana, N slope 8-VIII-6O (AOS).
77. Plebejus (Plebejus) saepiolus (Bdv.) Glacier 9- VII- 33 (JSG); Re-
7^77^~T7- Vn-33 (JSG); Florence 5-Vin-33 (JSG); Pate 23-Vn-34
(EDG); Tioga Pass 15-VIII-51, 6- VIII- 53 and 16 to 18- VIE- 57 (JWT),
9- VII to 24- VIII- 58 and 13 to 15- VII- 60 (AOS); Crane Flat 3- VII- 54
and 19- VII- 57 (JWT); Hetch-Hetchy Summit 13-VII-56 (JSG); Aspen
Valley 14- VII- 56 (JSG); Tuolumne Meadows 18- VII- 57 (JWT); Gin
Flat I9-VII-57 (JWT); Dana, W slope 17- VIII- 57 and 20- VII to 30-
Vin-58 (JWT); Crest W of Tioga Pass 9~Vn to 20-Vni-58 (AOS);
Helen Lake 18- VII- 58 and 19- VII- 60 (AOS); Rafferty Creek Trail
3-Vin-58 (JSG, AOS); Gaylor Lakes Trail 3 1- VHI to 3-IX- 58 (JWT);
Bridal Veil and Badger Pass 23- VI- 59 (JSG); White Wolf, Tioga Road,
and Crane Flat 24- VI-59 (JSG). Mono Lake 5-VII-38, 30- VI- 50 and
14-Vni-57 (JWT), 30-VI-41 (JSG), 13-VII-60 (AOS).
78a. Plebejus (Icaricia) icarioides icarioides (Bdv.) Glacier 9-VII-33
TjSG)r^si"en Valle7~T4- VII- 56 (JSGF” below Vogelsang Pass 3-Vni-
58 (AOS); Badger Pass 23- VI- 59 (JSG); Crane Flat 24- VI- 59 (JSG);
Jerseydale 29 to 30- V- 59 and 1 1- VI- 6 1 (JWT), 26- VI- 59 (JSG), 9
to 13-VI-62 (AOS); Indian Flat 13- V-6 1 to 10- VI-6 1 (JWT); Mather
10- VI-61 and 30-VI-62 (JWT).
b. Plebejus (Icaricia) icarioides hellos Edw, Mono Lake d-VII-SS and
TJTvLhl (JWT), 37^-41 and 19-VI-54 (JSG), 7 to 20- VII- 58 and
26 to 27- VI-61 (AOS); Conway Pass, ‘ N of Mono Lake 30- VI-50 and
23-VI-52 (JWT); Huntoon Public Camp, Bridgeport 19- VI-54 (JSG);
Bodie 28- VII- 57 (AOS); Agnew Pass and Donohue Pass 4-Vni-59
(AOS); Warren Creek ll-VIII-bO (AOS); W of Lee Vining 26- VI- 61
(AOS); 5 mi N of Mono Lake 28- VI-61 (AOS).
Plebejus (Icaricia) shasta comstocki Fox Reserve l6-Vn-33 (JSG);
Kuna 7-Vn-33 (JSG); Ke rTI^ 2 7^ VII- 3 4 (EDG); Slide Creek 29- VE-
34 (EDG); Tioga Pass 13-VIII-50, 15-VEI-52, 5 to 6-VEI-53, and
16-Vin-57 (JWT), 15-Vn-57, 14-Vn to 20-Vm-58 and 15 to 20-VE-
86
GARTH AND TILDEN
/. Re$. Lepid.
60 (AOS); Pilot Peak 15- VIT 57 and 19- VH to 3 H VIII- 58 fJWT); Gin
Flat 17-Vni-57 (JWT); Dana, W slope 17-VIII-57 and 30-Vni-58
(JWT); Upper Gaylor Lake 14 to 19- VII- 58 (AOS); Helen Lake 18- VH-
58 (AOS); Upper Lyell Base Camp 9-VnL58 (AOS); Crest W of
Tioga Pass 19 to 20-Vni-58 (AOS); Dana, N slope 24-Vin-58 and 8-
VIII-60 (AOS); Gaylor Lakes Trail 3-IX-58 (JWT). Bodie 18-VIL
57 (AOS); Lundy ZO-VII-58 (AOS); Donohue Pass 4 to 6~ VIII- 59
(AOS); Warren Creek ll-Vni-60 (AOS).
80, Plebejus (Icaricia) acmon (West. & Hew.) Reserve 14-VII-33 (JSG);
pShonT“ 25^ra33 "(JS^ Kerrick 27-VII-34 (EDG); Coldwater 3-
Vm-34 (EDG); Smoky Jack 4-Vn-54 (JWT); Mather lltol3-VII-
56 (JSG); Hetch-Hetchy Stimmit 13-VII-56 (JSG); Gin Flat IT'-VIII-
57 (JWT); Tenaya Canyon ll-VII-SS (AOS); Tuolumne Meadows 3-
.IX-58 (JWT); Jerseydale 30-V-59, 27-V-60 and ll-VI-61 (JWT),
26- VI-59 (JSG); Indian Flat 30-V-59, 27-V-60and 14-V to 10-VI-
61 (JWT); Briceburg 21-111-61 (KT); El Portal 21-111-61 (KT), 14-
V-61 (JWT); Cliff House 9-VI-61 (JWT); Mather 10- VI-61 (JWT);
Cathay ll-VI-61 (JWT). Mono Lake 30- VI-41 (JSG), 30- VI to 7-
Vn-58 and 12- VI- 62 (AOS); Huntoon Public Camp, Bridgeport 19- VI-
54 (JSG); Warren Creek ll-VIII-60 and 26- VI-61 (AOS).
81, Plebejus (Icaricia) lupini (Bdv. ) Eagle Peak l-Vn-33 (JSG); Dana,
W slope l6^VnL52T'T^VIII-53 and 20-Vn-58 (JWT); Smoky Jack 3
to 4- VII- 54 (JWT); Crest W of Tioga Pass 12-VI1-57, 4-Vn to 19-
Vni-58, 15 to 23-Vn-60 and 25-VI-61 (AOS); Upper Gaylor Lake 14-
to 19-VII-58 (AOS); Helen Lake 18- VII- 58 (AOS); Dana, N slope 24-
Vm-58 (AOS), Lee Vining 3 to 5-Vn-38 (JWT); E above Saddlebag
Lake 28-VII-58 (AOS); Below Conway Summit 23-VI-62 (JWT).
82, Everes comyntas (Godt. ) Mono Lake 5-Vn-38, 30- VI- 50 and 23- VL
6TTJW T)r~l6^I- 6 1 and 12- VI- 62 (AOS); E above Saddlebag Lake
28-VII-58 (AOS),
83, Everes amyntula Bdv. Hetch- Hetchy Siimmit 13-VU-56 (JSG, seen);
Jerseydale 30^-59 (JWT).
84, Philotes enoptes (Bdv.) Yosemite Valley 17-VI-32 (JWT); Ledg-e 9-
VIL33 IjSOF^eserve l6-Vn-33 and 7-Vn-56 (JSG); Tioga Pass
16-VIL52 (JWT), 12-Vn-57, 9-Vn to 14-Vni-58 and 14 to 23-Vn-60
(AOS); Smoky Jack 4- VII- 54 (JWT); Big Oak Flat Road 5- VII- 54
(JWT); Tenaya Canyon ll-VII-58 (AOS). E above Saddlebag Lake
28- ¥11- 58 (AOS).
85a. Philotes battoides battoides (Behr) Reserve 15 to 16-VII~33 and 7-
Vn-56 (JSG);“’^dr'c?^‘^^ 29-Vn-34 (EDG); Gaylor Lakes Trail 16-
Vni-52, 6-Vni-53 and 3.IX-58 (JWT); Mt. Hoffman 14.Vn-57 (AOS);
Pilot Peak 18- Vn-57 and 3 1- Vin-58 (JWT); Crest W of Tioga Pass
19~Vn to l-Ym-58, 14 lo 15-Vn-60, 25- VI- 61 and 8- Vm- 62 (AOS);
Helen Lake 18to29“Vn-58 (AOS); Cockscomb Peak 26-VII-58 (AOS);
Vogelsang 23-Vn-58 (JWT); Vogelsang Pass 3- ¥111-58 (AOS),
b. Philotes battoides glaucon (Bdv.) Mono Lake 5- VII- 38 and 23- VI- 62
(JWT), 30-YI-41 NSGfr^ 30- VI-58 and 26 to 27-VI-61 (AOS); War-
ren Creek 21-Vn~58 (AOS); Conway Summit 23-VI-62 (JWT).
86, Philotes speciosa (Hy, Edw. ) Mariposa 30-V-32 (G. & Bohart);
(R. M, Bohart),
87, Phaedrotes piasus (Bdv.) Eagle Peak l-Vn-33 (JSG); Reserve 9 to
TATvn^i^lED^; Jerseydale 9 to 13- VI- 62 (AOS); Mather 30- YI-
62 (JWT). Mono Lake 30-VI-41 (JSG).
2(i):i-96, 1963
YOSEMITE BUTTERFLIES
87
88a. Glaucopsyche lygdamus behrii(Edw.) Indian Flat 16 to 17-IV-61 (JWT);
El Porting- IV- 6F7JwT)7“
b. Glaucopsyche lygdamus Columbia (Skin.) Eagle Peak l-VII-33 (JSG);
Reserve 9 to 14- VII-34“[eDG ?); Tioga Pass 16- VIII- 52 (JWT);
Crane Flat and Tamarack Flat 3- VII- 54 (JWT); White Wolf 10- VII-
56 (JSG); Tenaya Canyon ll-Vn-58 (AOS); Upper Gaylor Lake 19-
VII-58 (JWT); Dana, W slope 20-VII-58 (JWT); Below Vogelsang
Pass 3-VIIL58 (AOS); Badger Pass 23-VI-59 (JSG). E above
Saddlebag Lake 28- VII- 58 (AOS).
89. Celastrina argiolus echo (Edw. ) El Portal 28- VI-33 (JSG); Glacier
'9- VII- 33 (JSG); ReT^e 14 to 18- Vn-33 and 7- VII-56 (JSG); Crane
Flat 3-VII-54 (JWT); Mather 12-Vn-56 (JSG), 14-V-61 (JWT);
Aspen Valley 14- VII- 56 (JSG); Darrah, Jerseydale, and Indian Flat
29 to 30-V-59 and 10- VI-61 (JWT); Badger Pass and Bridal Veil 23-
VI- 59 (JSG); Tamarack 24- VI- 59 (JSG); Briceburg and El Portal
21-III-61 (KT).
PIERIDAE
90a. Anthocharis (Anthocharis) sara sara Luc. Indian Flat 13-V-61 (JWT);
El Portal 14-V-61 (Jl^). “ “
Anthocharis (Anthocharis) sara reakirtii Edw. , gen. vern. Brice-
burg 21-III-6T~(W), 16- VI- 61 (JWT); El PortST" 21-III and 4-IV-
61 (KT), 14-IV-62 (JWT); Indian Flat 17-IV-61 (JWT).
b. Anthocharis (Anthocharis) sara stella Edw. 'Eagle Peak l-VII-33
*^S^ ReserT^ 16 to 17- VII-33" ( JSG);"'"- White-'Wolf lO-Vn-56 (JSG).
91. Anthocharis (Falcapica) lanceolata Luc. Eagle Peak l-VII-33 (JSG);
Ledge 9 - VII- 3 3 (JSG); Tosemltr Valley 14-V-59 (R. Patil Allen);
Jerseydale 9 to 13-VI-62 (AOS).
92. Euchloe creusa hyantis (Edw.) Eagle Peak l-VII-33 (JSG); Glacier
9^VILT3 (JSG); Reserve l6-Vn-33 (JSG); Ostrander 4- VII- 34
(EDG); El Portal 16- IV and 14- V- 6 1, andl4-IV-62 (JWT, common);
Indian Flat 17-IV-61 (JWT); Mather 14-V-61 (JWT).
93. Euchloe ausonides coloradensis (Hy. Edw.) Reserve 16 and 17-VII-
33 TJSG); Tamara^ Flat 3- VII- 54 (JWT). Mont) Lake 30- VI- 58
(AOS).
94. Colias eurytheme Bdv. Ledge 9- VII- 33 (JSG); Reserve l6-Vn-33
and 7 to“TlTviI-56 (JSG); Camp 9, Yosemite Valley 23-Vn-33 (JSG);
Crane Flat 3-VII-54, 18-Vm-57, and 10- VI-61 (JWT); Mather 11
to 12-VII-56 (JSG), 14- V and 10- VI- 61 (JWT); Hetch-Hetchy 13-VII-
56 (JSG); Aspen Valley 14- VII- 56 (JSG); Gaylor Lakes Trail 19- VII-
58 (JWT); Dana, W slope 30-VIII-58 (JWT); Briceburg 21-III-61
(KT); El Portal 21-III-61 (KT), 14.V-61 (JWT); Yosemite Valley
I9-IV-6I (KT); Indian Flat 13-V-61 (JWT); Jerseydale II-VI-6I
(JWT).
95. Colias philodice hagenii Edw. Mono Lake 30-VI-41 (JSG); 18-VII-
57 and 30-VI-58 "JKosY.
96. Colias occidentalis chrysomelas Hy. Edw. "Yosemite" VI-26 (E.
O. Essig).
Colias behrii Edw. Tioga Pass 13-Vni-29 and 1 to 3-Vin-56 (JSG),
14-VII-31, 13-Vin-50 and l6-Vni-57 (JWT), 4-Vni-57 and 14-Vnto
19-Vni-58 (AOS); Florence 5-Vni-33 (JSG); Kuna 7-Vin-33 (JSG);
Kerrick 27-VII-34 (EDG); Snow Lake 29-Vn-34 (EDG); Tuolumne
88
GARTH AND TILDEN
/. Res. Lepid.
Meadows 15-Vni-57 (JWT); Pilot Peak 15 to 18- VIU- 57 (JWT);
Upper Gaylor Lake 19-VII to 20-Vni-58 (AOS); Helen Lake 29-VII-
58 (AOS); Crest W of Tioga Pass 31-VII-58 (AOS); Rafferty Creek
Trail 3-Vni-58 (AOS, JSG); Merced Lake Trail 4-VIII-58 (AOS);
Lyell Base Camp 10- VIII- 58 (AOS); Elizabeth Lake 23- VIII- 58 (AOS).
E of Donohue Pass 4 to 6-VIII-59 (AOS),
98. Colias (Zerene) eurydice Bdv. "Yosemite" VI-26 (E, O. Essig or
PTJTWooIfJT"^
99. Nathalis iole Bdv. "Yosemite" VI-26 (P. J. Woolf). Warren Creek
20, 21 ai^n8-VU-58 (AOS).
100. Neophasia menapia tau (Scud.) Big Oak Flat Road VIII-28 (JSG, seen);
Little Yosemite ValT^ 3- VIII- 33 (JSG); Pate 23- VII- 34 (EDG);
Buck Meadow 4-VII-54 (JWT); Unicorn Peak 27-VII-59 (AOS). Dead-
man's Summit 20-IX-56 (JWT); June Lake 2 1-VIII-57 (JSG),
101. Pieris (Pontia) beckeri Edw. Mono Lake 5-VII-38, 30-VI-50, 15-
Vni-53 and 23 -VI- 62 (JWT), 26 and 27-VI-61 (AOS).
102. Pieris (Pontia) sisymbrii Bdv. Eagle Peak l-VII-33 (JSG); Reserve
16- VII- 33 and 7-Vn-56 (JSG); Dana, W slope 19"Vn-52, 16-VIII-53
and 20-Vn-58 (JWT); Upper Gaylor Lake 4 to 8-VII-57, 9 to 19-Vn-
58 and 25-VI-61 (AOS); Crest W of Tioga Pass 9-VII-58 (AOS);
Tioga Pass lO-Vn-58 (AOS); Briceburg and El Portal 2 l-ni-6 1 (KT);
El Portal 14-IV-62 (JWT).
103. Pieris (Pontia) protodice Bdv. h Lee. El Portal 28-VII-33 (JSG);
Yosemite Creek Trail 9 to 17-Vn-56 (JSG).
104a. Pieris (Pontia) occidentalis occidentalis Reak. Crest W of Tioga Pass
181^20- Vm- 57 and 19 to 2”0-VIII-58 (AOS),
b. Pieris (Pontia) occidentalis calyce Edw. Pilot Peak 18- VIII- 52, 15-
Vni-57, and“20-VII-58 (JWT); Crest W of Tioga Pass 8-VII-57 and
9-VII-58 (AOS); Upper Gaylor Lake 9-VII-58 (AOS), l-Vm-58
(JSG); Vogelsang Camp 3-VIII-58 (JSG). Warren Creek 29-VIII-57
(AOS).
105. Pieris (Pieris) napi venosa Scud. Yosemite Valley 14-V-59 (R. P.
Allen); Briceburg 21-III-61 (KT); El Portal 14-IV-61 (JWT); Jer-
seydale 22-VI-61 (AOS).
106. Pieris (Pieris) rapae (Linn.) Pate 23-VII-34 (EDG); Indian Flat 30-
V-59 (JWT, se^l7“
PAPILIONIDAE
107. Papilio zelicaon Luc. Reserve l6-Vn-33 (JSG); Mather 15-VII-56
(JSG); Jerseydale 16- VI- 57 and 4- VUI- 62 (AOS); Crest W of Tioga
Pass 9-VII-58 (AOS); Upper Gaylor Lake 19-VII-58 (AOS); El Por-
tal 4-IV-61 (KT).
108. Papilio indra Reak, Yosemite Valley 17-VI-32 (JWT); Reserve 18-
VII- 3r (A. Carthew); Crest W. of Tioga Pass 25- VI-61 (AOS).
Mono Lake 23-VI-62 (JWT). Mammoth Peak 16-VII-60 (AOS).
109. Papilio rutulus Luc. Camp 19, Yosemite Valley 26- VI-33 (JSG);
Pate 23- VII- 34 (EDG); Yosemite Creek Trail 9-Vn-56 (JSG);
Hetch-Hetchy 13-VII-56 (JSG); Darrah, Jerseydale, and Indian Flat
29 to 30-V-59 (JWT). Mono Lake 30- VI-41 (JSG, seen), 27-VI-61
and 12- VI- 62 (AOS); W of Lee Vining 27-VI-61 (AOS); Below Conway
Summit 23- VI-62 (JWT).
Papilio multicaudatus Kby. Jerseydale 7 -VIII- 5 6 and 2- VIII- 57 (AOS).
2(i)ti-s6, 196}
YOSEMITE BUTTERFLIES
89
Ilia. Papilio eurymedon eurymedon Luc. Ledge 9-VII-33 (JSG); Pate 23-
Vn-34" (EDG); Mather 12-^-56 (JSG); Hetch-Hetchy 13-Vn-56
(JSG); Jerseydale 16- VI- 57, 16- VI-58, 21-VI-61 and 13-VI-62 (AOS);
Darrah, Jerseydale, and Indian Flat 29 to 30-V-59 (JWT).
b. Papilio eurymedon albanus F. & F. Research Reserve 10- VII- 56
IjSGir
112. Parnassius clodius baldur Edw. Tioga Pass 7-VII-31 (JWT); Eagle
Peak 1-VII^3 (JSG)r^lacier 9- VII- 33 (JSG); Dana 8- Vni-33 ( JSG);
Slide 29-VIL34 (EDG); Gin Flat 4-Vn-54 (JWT, seen); Jerseydale
23-VI-56, 21-VI-57. 16 to 18- VI- 58 and 22- VI- 6 1 (AOS); Yosemite
Creek 7-Vn-56 (JSG); White Wolf 10- VII- 56 (JSG); Crest W of
Tioga Pass 12-VII to IS-VIH-ST, 28- VII to 19-VIII-58 and 4 to 20-VII-
60 (AOS); Pilot Peak 18- VII- 57 and 31-Vin-58 (JWT); Rafferty Creek
21- VII- 57 (AOS); Tenaya Canyon ll-VII-58 (AOS); Lewis Creek
Trail 3-Vni-58 (JSG); Dana, W slope 30-VIII-58 (JW-T).
113. Parnassius phoebus behrii Edw.
Reserve 14- VH^S' (“180)7 Dana8-Vm-33 (JSG); Pilot Peak 13 to
14- VIII- 52, 18-Vn to 15-Vni-57 a-nd 31-VIII-58 (JWT); Upper Gaylor
Lake 3 1- VII to 20- VHI- 57 and 3 1- VII to 20- VIU- 58 (AOS), 19-VII-58
(JWT), l-Vin-58 (JSG); Helen Lake 18- VII- 58 (AOS); Dana, W
slope 20-VII-58 (JWT); Crest W of Tioga Pass 19-VIII-58 (AOS);
Dana, N slope 24- VIE- 58 (AOS). E of Saddlebag Lake 28-VII-58
(AOS). .
114. Epargyreus clarus (Cram j Coulterville Road VI-26 (D. D. MacLean);
Indian Flat 30-V-59 (JWT, seen).
115. Thorybes pylades (Scud.) Fish Camp 18- VI- 57 (JWT); Jerseydale
JO^V^T^ (“jWrjT^Indian Flat 13-V-61 (JWT).
116. Thorybes mexicana nevada Scud. Tioga Pass 7-VII-31, l6-Vni-52
and 18-VIL5TnJ^Tir~8TviI-57, 26- VI to 19- VH- 58, 14-VII-60 and
25-VI-61 (AOS); Reserve 16-VII-33 (JSG); Tamarack 3-Vn-54
(JWT); Pilot Peak 18-VII-57 and 3 1- VIE- 58 (JWT); Dana, W slope
I7.VEI-57 and 20-VE-58 (JWT); Gaylor Lakes Trail 9-VII-58 (JWT);
Crest W of Tioga Pass 9-VE-58 (AOS); Upper Gaylor Lake 14 to
19-VII-58 (AOS), l-VIE-58 (JSG); Helen Lake 18 to 29-VII-58 (AOS);
Gin Flat and Crane Flat lO-VI-61 (JWT).
117. Thorybes diversus Bell Aspen 14-VE-56 (JSG); Jerseydale I6-VI-
5Ta^n6-"vi-58 fAOS); Mather 10- VI- 61 and 30- VI- 62 (JWT).
118. Pyrgus ruralis (Bdv. ) Reserve 15- VE- 33 and 10- VII- 56 (JSG); Tam-
arack 3-VII-54 (JWT); Tenaya ll-VII-58 (AOS, seen); Lyell Fork
Meadows 7- VIE- 58 (AOS).
119. Pyrgus communis (Grote) "Yosemite" VI-26 (P, J. Woolf); Aspen
' Valli^ 14-VE-56" (JSG); El Portal 16-IV, 14-V and lO-VI-61 (JWT);
Indian Flat 15-IV, 14- V and 1 1- VI-61 (JWT),
120. Heliopetes ericetorum (Bdv.) Jerseydale 16-VI-58 (AOS); El Portal
14- V-61 (JWT, s7^n)7 Highway 120 and Hetch-Hetchy Road 14-V- 61
(JWT).
121. Erynnis persius (Scud.) Reserve 15-VII-33 (JSG, as afranius); Chow-
chilla Mt. 26-VI-54 (JWT); Aspen Valley 14-VII-56 (JSG); Mather
15- VE-56 (JSG); Badger Pass 23-VI-59 (JSG); Indian Flat 17 -IV- 6 1
(JWT); Jerseydale 21to23-VI-6l (AOS).
122. Erynnis pacuvius lilius (Dyar) Chowchilla Mt. 26- VI- 54 (JWT, com-
mon); Tamarack fET™ 3 - VII- 54 (JWT); Crane Flat 24- VI- 59 (JSG).
Erynnis propertius (Scud. & Burg.) Ledge 9-VII-33 (JSG); Reserve
90
GARTH AND TILDEN
/. Res. Lepid.
16-VII-33 (JSG); Crane Flat 3-Vn-54 (JWT); Yosemite Creek 9-VII-
56 (JSG); Mather 12-VII~56 (JSG), 10- VI- 61 (JWT); Hetch-Hetchy
Sixmmit 13-Vn-56 (JSG); Badger Pass 23-VI-59 (JSG); Briceburg
21-in-6l (KT); El Portal 21-ni-6l (KT), 14-V-61 (JWT); Yose-
mite Valley 19-IV-61 (KT); Indian Flat 13-V and 10- VI-61 (JWT);
Jerseydale 17- VI- 60, 2 1- VI- 6 1 and 4 to 13- VI- 62 (AOS).
Erynnis zarucco fnneralis (Scud. & Burg.) Coulterville Road VI-26
(D. D. MacLean).
125. Erynnis tristis (Bdv. ) No Yoserhite record.
126a. Hesperia harpalus harpalus (Edw. ) Mono Lake 17-VII to 26-Vni-57,
30-VI to 20-VII-58 and 26-VI-61 (AOS), 26-Vin-57 and 20- VH- 58
(JWT); Bodie 18-VII-57 (AOS); Lee Vining 24- VI-58 (AOS); War-
ren Creek 26-VI-61 (AOS).
Hesperia harpalus (Edw.), "blend zone" form Eagle Peak l-VII-33
(JSG); Reserve 16- VII- 33 (JSG); Yosemite Creek 9 to 17-VII-56
(JSG); Rafferty Creek 21-Vn-57, 24- Vn to 3- Vin-58 (AOS); Dana,
W slope 16-VIII-57 and 30-VIII-58 (JWT); Crest W of Tioga Pass 18
to 20-VIII-57, 20-Vin-58 and 20 to 28-Vn-60 (AOS); Merc.ed Lake
Trail 4-VIII-58 (AOS); Unicorn Peak 12- VIII- 58 and 27- VB- 59 (AOS);
Gaylor Lakes Trail and Pilot Peak 31-VIII-58 (JWT); Tuolumne
Meadows 3-IX-58 (JWT); Cold Canyon 25-Vn-60 (AOS); Dana, N
slope 8-Vin-60 (AOS).
b. Hesperia harpalus yosemite Leuss. Big Oak Flat Road, near entrance
to Tuolumne Grove 2- IX- 29 (J. Strohbeen and A. E. Dodge, the type
series); Carl Inn ll-VII-31 (JWT); Wawona 27-Vm-46 (JWT);
Smoky Jack 4-VII-54 (JWT); Jerseydale l-Vin-57 (AOS).
127. Hesperia miriamae MacNeill Mono Pass, Inyo County VIII- 56, VIII-
57 and Vni-58 (Don MacNeill, the type series); Unicorn Peak 26-Vn,
12 and 23- VIII-58, and 27-Vn-59 (AOS); Cockscomb Peak 26-Vn-58
(AOS); Dana, N slope 31- VII- 58 and 8-Vin-60 (AOS).
128. Hesperia nevada (Scud.) Mono Lake 26-VI-61 (AOS); Warren Creek
26- VI- 61 and 12"-VI-62 (AOS).
129. Hesperia juba (Scud.) "Yosemite" VI-26 (collector unknown); Old
Village lTrvj-32 (McQuesten); Mather 10- VI-61 (JWT). Mono
Lake 27- VI-61 (AOS).
130. Ochlodes sylvanoides (Bdv.) Yosemite Valley Vn-33 (JSG); Pate
Van^7~23”vil- 34 TeDG); Hetch-Hetchy 13- VII- 56 (JSG); Crane
Flat 18- VIII- 57 (JWT); Mather 4-IX-57 (JWT); Jerseydale 2-IX-
58 (AOS). Mono Lake 14 and 15-Vm-51 and 13 to 15-VIII-52 (JWT).
13 1. Ochlodes agricola (Bdv. ) Darrah 29-V-59 (JWT); Indian Flat 30-V-
59 (JWT); El Portal 14-V-61 (JWT).
132a. Polites sabxileti sabuleti (Bdv.) Bridgeport 14-Vni-50 and 16-Vni-51
(JWT); Mono La'ki^2“3rvi- 62 (JWT).
b. Polites sabuleti tecumseh Grin. Tioga Pass 7-Vn-31, 13-VIII-50,
Ts^^^-TITTs^VLElSZ and 18-Vnto 16- VIII- 57 (JWT), 14-VIIto7-
VIII- 58 (AOS); Reserve 17-Vn-33 and 10- VII- 56 (JSG); Kuna 7-Vm-
33 (JSG); Kerrick 27-VII-34 (EDG); Snow Lake 29-VII-34 (EDG);
Tuolumne Meadows 15-VIII-57 and 3-IX-58 (JWT), 15-Vin-58 (AOS);
Gin Flat 17-Vn-57 (JWT); Rafferty Creek 24-VII-58 (AOS), 3-Vin-
58 (JSG); Upper Gaylor Lake l-Vin-58 (JSG); Upper Lyell Base
Camp 9-Vin-58 (AOS); Elizabeth Lak% 12-Vin-58 (AOS); Dana, W
slope 30-Vni-58 (JWT); Pilot Peak 31-Vin-58 (JWT).
2(t)tl-96, 196}
YOSEMITE BUTTERFLIES
91
133. Polites sonora (Scud, ) Ledge 9-VII-33 (JSG); Reserve 17-Vn-33
(JSG); Tioga Pass l6-Vni-52 (JWT); Aspen Valley 14-Vn-56 (JSG);
Crane Flat 18 and 19"Vn-57 (JWT); Gin Flat 19-VII-57 (JWT);
Tuoltimne Meadows 14-Vin-57 and 3-IX-58 (JWT), 15 to 21-VIII-58
(AOS); Bridal Veil 23-VI-59 (JSG); Mather 10- VI- 6 1 and 30- VI- 62
(JWT). Mono Lake 4-VII-33, 5-VII-38, 30-VI-50 to IS-VIII-SO and
15-Vni-51 (JWT), 15-Vni-57, 30-VI-58 and 27-VI-61 (AOS); Bodie
18- VII- 57 (AOS); Warren Creek ll-VIII-60 (AOS); W of Lee Vining
26-VI-61 (AOS).
134. Amblyscirtes vialis (Edw.) Yosemite Valley, Meadow near Ahwahnee
HStirT7^VL3T'‘7jWT); Jerseydale 17- VI- 57, 22- VI- 6 1 and 9 to 13-
VI- 62 (AOS).
YOSEMITE PARK
1933 Localities: 1 — Camp 19; 2 — ^Camp 9; 3 — Mt. Dana; 4 — Eagle Pk.;
5 — El Portal; 6 — Florence L.; 7- — Glacier Pt.; 8 — Glen Aulin; 9 — 'Mt.
Kuna; 10 — Ledge Tr.; 11 — Little Yosemite; 12 — Mt. Lyell; 13 — Mu-
seum; 14 — Pohono Tr.; 15 — Research Reserve; 16 — Wawona.
1934 Localities: 17 — Benson L.; 18- — Coldwater Can.; 19 — KerrickPass;
20 — Ostrander L.; 21 — Pate Valley; 22 — Slide Can.; 23 — Tenaya L.
Key
Auto Rd.
- - Trails Covered 1933 .... Trails Covered 1934
92
GARTH AND TILDEN
/. Res. Lepid.
APPENDIX III
YOSEMITE LOCALITIES
West- slope localities visited by John S. Garth in 1933 (See Map 1):
1. Camp 19; Yosemite School of Field Natural History headquarters near
Sentinel Bridge, floor of Yosemite Valley (3960). Transition Zone;
Meadow Association.
2. Camp 9: Meadow near Camp Curry in Yosemite Valley (4000), a fav-
orite haunt of Argynnis cybele leto. Transition Zone; Black Cotton-
wood; Meadow Association.
3. Dana: Trail from Tuolumne Meadows (8600) to top of Mt. Dana (13, 050).
Hudsonian, Arctic-Alpine Zone; Subalpine Forest; Alpine Fell-Fields.
4. Eagle: Trail from top of Yosemite Falls (6525) to summit of Eagle Pk.
Canadian Zone; Red Fir Forest; Moist Meadow; Riparian Association.
5. El Portal: Highway 140 near El Portal (1919) on the Merced River; short
distance up Coulterville Road. Upper Sonoran Zone; Chaparral; Stream.
6. Florence: Lake above Lewis Creek Trail (10,500). Hudsonian Zone, in
close proximity to Arctic- Alpine. Subalpine Forest; Montane Meadows.
7. Glacier Trail from Glacier Point (7214) to top of Nevada Falls (5910).
Canadian Fir Forest replaced by Upper Sonoran Chaparral after fire.
8. Glen Aulin: High Sierra Camp on Tuolumne River at confluence of Con-
ness Creek (7850). Canadian Zone; Transition elements (A. hydaspe).
9. Kuna; From Lyell Base Camp (10, 600) to summit of Kuna Crest (12, 000),
down Dana Fork to Tuolumne Meadows (8600). Hudsonian, Arctic-Al-
pine Zone; cirques, tarns, snow banks, talus slopes, alpine meadows.
10. Ledge: Short trail from Camp Curry (4000) to Glacier Point (7214).
Transition, Canadian Zone; steep cliffs, talus slopes, springs, stream,
11. Little Yosemite: Hanging Valley above Nevada Falls (5910) on Merced
Lake Trail. Transition Zone; Jeffrey Pine, White Fir, Western Juniper.
12. Lyell: From Vogelsang Lake (10, 500) to summit of Mt. Lyell (13, 090).
Hudsonian, Arctic- Alpine Zone; glaciers, tarns, Alpine rock gardens.
13. Museum: Garden along diverted stream behind Yosemite Museum (4000).
14. Pohono: Trail from Glacier Point (7214) over Sentinel Dome (8117) to In-
spiration Point (5391). Canadian and Transition Zone; stream, meadow.
15. Reserve; Boundary Hill Research Reserve, a 25- square-mile area bound-
ed by the Tioga Road, Yosemite Creek, Cascade Creek, and the north
rim of Yosemite Valley. (6525 - 9200). Canadian Zone with weak Hud-
sonian atop Research Ridge; Red Fir Forest; Lodgepole Pine; Western
Juniper.
16. Wawona: Highway 41 (Wawona Road) from Chinquapin (6266) to Wawona
(4096). Transition Zone; Dry Hillside; Monardella and Erysimum.
West- slope localities visited by Edmund D, Godwin in 1934 (See Map 1):
17. Benson: Trail from Pate Valley (4500) over Pleasant Ridge (8000) to
Pleasant Valley (7000) and Benson Lake (8000). Transition, Canadian
Zone; Upper Sonoran influence (Cercyonis silvestris, Coenonympha).
18. Coldwater: Trail from Virginia Creek (8500) to Glen Aulin (7800) along
Coldwater Canyon. Canadian Zone; Red Fir Forest; Riparian Assoc.
19. Kerrick: Trail from Benson Lake (8000) through Seavey Pass (9100) to
Kerrick Canyon (8900), thence to Kerrick Meadows (9400). Canadian
to Hudsonian Zone; Red Fir, Lodgepole Pine Forest; Montane Meadow.
2(i)xi-96, 196}
YOSEMITE BUTTERFLIES
93
20. Ostrander: Trail from Glacier Point Road (7100) to Ostrander Lake
(8500). Canadian to Hudsonian Zone border (Tsuga mertensiana).
21. Pate: Trail from Harden Lake (7575) down north- facing granite slope
to Pate Valley (4500). Canadian to Transition Zone; Upper Sonoran
influence from opposite exposure. Monardella and Umbellularia.
22. Slide: Trail from Kerrick Meadows (9400) to Snow Lake (10, 200) to
Slide Canyon (10, 000). Hudsonian Zone; Subalpine Meadow Associa-
tion (Colias behrii).
23. Tenaya: McGee Lake Trail from Glen Aulin (7850) on the Tuoltimne
River to Tenaya Lake (8141). Canadian Zone; Red Fir; Lodgepole Pine.
Western Foothill localities visited from 1956 to 1962, mostly outside
Yosemite National Park:
Bear Creek Lodge: On Highway 140 between Midpines (2477) and Brice-
burg(1186). Transition Zone; Yellow Pine Forest; Riparian Association.
25. Big Oak Flat Road: Highway 120, the part between Groveland (2846) and
Carlon (4624) within Stanislaus N F. Upper Sonoran, Transition Zone.
26. Briceburg: Mariposa Co. , on Highway 140 in Merced River Bottom
(1186). Upper Sonoran Zone; Chaparral and Coastal Sage Scrub.
27. Buck Meadows: Stanislaus N F, on Highway 120 (3006). Transition Zone;
Yellow Pine Forest with "Mountain Misery" (Chamaebatia) understory.
28. Carl Inn: Stanislaus N F, at juncture of Hetch-Hetchy Road to Mather
and Old Tioga Road (4718). Transition Zone; Mixed Forest, cut over.
29. Cathay: Mariposa Co., on Highway 140; Upper Sonoran; Oak Woodland.
30. Chowchilla Mtn. : Sierra N F, W of Fish Camp (4982) on Chowchilla Mtn.
Road, near Bear Wallow turnoff. Canadian Zone; Red Fir Forest.
31. Cliff House: Stanislaus N F, on Highway 120 N Fork, Tuolxxmne River,
crossing. Transition Zone; Yellow Pine Forest; Y erba Santa.
32. Darrah: Mariposa Co. (3400). Upper Sonoran, Transition Zone; Chap-
arral; Coastal Sage Scrub; Yellow Pine, Mixed Forest; Riparian Assoc.
33. El Portal: Yosemite N P, on Highway 140 in Merced River Bottom (1919).
Upper Sonoran Zone; Chaparral, Coastal Sage Scrub; Riparian Assoc.
34. Groveland; Stanislaus N F, on Highway 120 (2846). Transition and Upper
Sonoran Zone; Yellow Pine Forest; Mixed Forest; Chaparral.
Highway 120 and Hetch- Hetchy Road: Juncture, Big Oak Flat Road and
Mather Road (4624). Upper Sonoran, Transition; Oak Wood; Yellow Pine.
36. Indian Flat; Sierra N F, on Highway 140 in the Merced River bottom
(1553). Upper Sonoran Zone; Coastal Sage Scrub, Chaparral; Yellow Pine.
37. Jer seydale: Sierra N F, (3800). Transition Zone; Yellow Pine, Mixed
Forest; Chamaebatia, Monardella (Argynnis hydaspe); some Meadowland.
38. Mariposa: Mariposa Co. , on Highway 140 (2046). Upper Sonoran, Transi-
tion Zone; Chaparral, Coastal Sage Scrub; Mixed Forest; Riparian Assoc.
Mather: Stanislaus N F (4522), on Hetch-Hetchy Road. Transition Zone;
Yellow Pine, Mixed and White Fir Forest; Montane Meadow; Riparian.
Includes Carnegie Experimental Garden (Hog Ranch) 1 mi NE of Mather.
40. Oakhurst: Madera Co., on Highway 41. Upper Sonoran Zone; Foothill
Woodland; Oak and Digger Pine with Ceanothus and grass (Coenonympha).
West-Slope Mid- Elevation Localities visited from 1956 to 1962, all within
Yosemite National Park:
41. Aspen Valley: On Old Tioga Road above turnoff from Mather Road at Carl
Inn (639O). Canadian Zone; Red Fir Forest; Montane Meadow; lupine.
94
GARTH AND TILDEN
/. Res. Lepid.
42. Badger Pass: Meadow beside Badger Pass Ski Lodge on Glacier Point
Road (7300). Canadian Zone; Red Fir Forest; Wet Meadow Association.
43. Bridal Veil: Public Camp on Bridal Veil Creek where crossed by Glac-
ier Point Road (7000). Canadian Zone; Red Fir Forest; Riparian Assoc.
44. Crane Flat: On Highway 120 where it becomes the Tioga Road (6192).
Transition Zone; Mixed Forest; Montane Meadow. Also roadside above.
45. Gin Flat: On Tioga Road above Crane Flat (7036), Canadian Zone;
Montane Meadow.
46. Glacier Point: Road to, with maximum elevation 7800 feet, Canadian
Zone; openings in Red Fir Forest; Montane Meadow; Riparian Assoc.
47. Hetch-Hetchy: Road from Mather (4522) to Hetch-Hetchy Reservoir
(3796). Upper Sonoran Zone; Digger Pine; Manzanita. Summit (5027)
Transition Zone; White Fir Forest; Streamside Association; lupine.
48. Mirror Lake: Floor of Yosemite Valley (4082). Transition Zone; Black
Oak, White Fir, Incense Cedar, Cottonwood, Ceanothus, Coffee Berry.
49. Old Village: Yosemite Valley floor (3964). Transition Zone; Riparian.
50. Research Reserve: Meadow in Blue Jay Creek (7900). Canadian Zone;
Red Fir, Lodgepole Pine. Research Ridge (8700) Dry Hillside Assoc.
51. Smoky Jack; Public Camp on Tioga Road, above Gin Flat. Canadian
Zone; Red Fir Forest; Montane Chaparral.
52. Tamarack Flat: Meadow above Tamarack Public Camp on Tioga Road
(6390). Upper Canadian Zone; Red Fir; Lodgepole Pine; Ceanothus.
53. Tioga Road; North-facing slope 1 mi W of White Wolf turnoff (8000) and
first summit beyond White Wolf (8090). Canadian Zone; Red Fir Forest.
54. Tuolumne Grove: On Highway 120 W of Crane Flat (5500-6000). Tran-
sition Zone; Mixed Forest; Big Tree; Riparian Assoc. Polygonia zephyrus.
55. White Wolf; One mi N of Tioga Road (8090). Canadian Zone; Red Fir
Forest; Montane Meadow Assoc. Plebejus saepiolus, P. glandon podarce.
56. Wawona: Open forest between Wawona Lodge (4096) and the turnoff to
Giant Forest (4350). Transition Zone; Yellow Pine Forest.
57. Yosemite Creek: Trail S from Public Camp on Tioga Road (7200). Can-
adian Zone; Riparian and Wet Meadow; Western Juniper on drier slopes.
58. Yosemite Valley: Floor of Valley (4000). Transition Zone; Mixed Forest;
Riparian Association. Includes Meadow near Ahwahnee Hotel. A. vialis.
High Country localities visited since 1956 (those near Tioga Pass since
1929):
59. Agnew Pass: Inyo N F (9946). Hudsonian Zone; open sage slope.
60. Bert Lake: N slope of Mt. Maclure (11, 700); Arctic- Alpine; rock gardens.
61. Cockscomb Peak: Cathedral Range (11,000); Arctic-Alpine; White-Bark
Pine; gradual slope; Spraguea in gravelly areas. Hesperia miriamae.
Dana, N slope: (11,000- 13,050 ). Arctic-Alpine; rockslides, flowers,
grass. Lycaena p. hypophlaeas. Dana, W slope: From Tioga Pass (9941)
to timber line (11, 000). Hudsonian to Arctic- Alpine; Subalpine Forest;
Alpine Fell-Fields.
Dog Lake Trail: From Tuolumne Meadows (8600) to Dog Lake (9240). Hud-
sonian Zone; Subalpine Forest; talus slides; rock gardens. Lycaena cupreus.
Donohue Pass: Inyo N F (11, 100). Arctic- Alpine; meadows, exposed rock.
65. Elizabeth Lake: Cathedral Range (9500). Hudsonian Zone; Lodgepole Pine;
meadows, willows, heather. Lycaena mariposa, Polites s. tecumseh.
Gaylor Lakes: Above Tioga Pass Checking Sta. (10, 500). Arctic-Alpine;
Alpine Fell- Fields, Includes Upper Gaylor Lake. Melitaea d. malcolmi.
Glen Aulin: On Tuolumne River (7850). Canadian Zone; Jeffrey Pine;
arid slope.
2([);i-9€, 1963
YOSEMITE BUTTERFLIES
95
68. Helen Lake: E of Kuna Crest (10, 896). Arctic-Alpine; White-Bark Pine.
69. Hoffmann, Mt. : W of May Lake (10,921). As above; flowers in gravel.
70. Lambert Dome: N of Tuoltimne Meadows (9400). Hudsonian; forested.
71. Lyell Base Camps: Head of Lyell Fork (11,000), Arctic- Alpine; treeline,
72. Lyell Fork Meadows: (9000). Hudsonian; Lodgepole Pine; willow stream.
73. Ottoway Lake: Clark Range (8000). Canadian Zone; Red Fir Forest.
74. Mammoth Peak: N end, Kuna Crest (12,225), Arctic- Alpine; exposed.
75. Merced Lake Trail: From Merced Lake (7100) to Tuolumne Pass (10, 100)
via Fletcher Creek, Canadian to Hudsonian; Coniferous Forest; Meadows.
76. Mono Pass: Inyo N F (10599), S of Tioga Pass, between Mt. Gibbs and
Mt. Lewis. Arctic- Alpine; Alpine Fell-Fields, Hesperia miriamae.
77. Pilot Peak: NW above Tioga Pass via Gaylor Lakes Trail (11, 100), Hud-
sonian, Arctic- Alpine; Subalpine Forest, Alpine Fell- Fields, (See 81).
78. Rafferty Creek: (8600- 10, 100). Hudsonian; Lodgepole Pine; Riparian.
79. Saddlebag Lake: Inyo N F (10, 050). Upper Hudsonian and Arctic- Alpine;
few trees, willows, grasses, some rock. Includes E above (10,500).
80. Tioga Meadow: (9941). Montane Meadow E of Tioga Pass Checking Sta.
Tioga Pass: (9941). Hudsonian Zone; Subalpine Forest; Glacial Moraine.
81. Tioga Pass, N.W above: (10, 500- 11,000). Tree-line Association; rock
gardens. Parnassius p. behrii, Pieris o. calyce, Oeneis c. ivallda.
82. Tenaya Canyon: (4, 000-8,000, average 6,000), Canadian Zone; stream
bed choked with firs, aspen, willows; exposed slopes (Philotes enoptes).
83. Thousand Island Lake: Inyo N F (9850), Hudsonian; Lodgepole Pine; sage.
84. Tuolumne Meadows: On Tioga Road (8600). Hudsonian; Lodgepole Pine,
extensive meadows. Site of Lembert Homestead; metropolis of C. behrii.
85. Unicorn Peak: Cathedral Range (10, 849). Arctic- Alpine; boulder j\imble.
86. Vogelsang; Trail from Vogelsang Camp (10, 100) to Vogelsang Pass (10,400)
and below. Hudsonian, Arctic- Alpine; Subalpine Forest; Alpine Fell-Fields.
East- slope localities visited during survey; mostly Mono Basin; all out-
side Yosemite National Park:
87. Bodie: 11 mi E of Highway 395 (8369). Pinyon; Great Basin Sage; Meadow.
88. Bridgeport: On Highway 395 (6743). Willow Thicket; Riparian Association,
89. Casa Diablo Hot Springs; Inyo N F, on Highway 395 (7198). Transition;
Great Basin Sage Brush in Jeffrey Pine Forest; sparse Streamside Assoc.
Conway Pass: On Highway 395 N of Mono Lake, 200 ft. below Summit
(8136). Great Basin Scrub (Desert Peach, Chrysothamnus, Purshia);
Montane Meadow; Aspen Grove.
91. Deadman Summit: Inyo N F, on Highway 395 (7734 or 8168). Transition;
Jeffrey Pine Forest; Purshia, Artemesia, Chrysothamnus, Eriogonum.
92. Gull Lake: Inyo N F (7600), on June Lake circuit. Moist Montane Meadow;
Willow Thicket; Brownie Thistle (Cirsium aca.ulescens^ A. n. apacheana.
93. June Lake: Inyo N F (7716). Junction of Highway 395 and June Lake Road.
Transition; Jeffrey Pine Forest; Great Basin Sage, Neophasia menapia,
Lee Vi ning Creek: Inyo N F (7000). Above turnoff to last campground on
Tioga Pass Road. Dry Hillside; Manzanita; Ceanothus, Bitter Cherry.
95. Mono Lake-; (6419). Many associations in close proximity: Freshwater
Marsh, Salt Sink Margin, Willow Thicket, Aspen Grove, Moist Meadow.
Mammoth: Inyo N F (7900). On Highway'395, about 2-3 mi below. Tran-
sition; Jeffrey Pine Forest; Great Basin Sage Brush; Great Basin Scrub.
L'u^dy^ Inyo N F (7766), Sage, Aspen, Willow. Collecting to 8700 feet.
Warren Creek: Inyo N F (9000-9500). N fork of Lee Vining Creek above
Tioga Road crossing. Canadian Zone; willowed stream, sage-aspen slope.
96
GARTH AND TILDEN
/. Res. Lepid.
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©H THE LEPIJ©©FTERA
YOSEMITE BUTTERFLIES: An ecological survey of the
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f
f}
■/
,/
•«.
\ie
f
I
%
■ "■3"<
' • . ^ V
<.
,4$‘:
Volume 2
Number 2
September, 1963
a quarterly published at
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
edited by: WILLIAM HOVANITZ
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Journal of Research on the Lepidoptera
2(2):97425, 1%3
114© W. Orange Grme Ape., Arcadia, California, U.S.A>
© C&pyright iftfj
QUANTITATIVE ANALYSIS OF CERTAIN WING AND
GENITALIA CHARACTERS OF PIERIS IN
WESTERN NORTH AMERICA
VINCENT C. S. CHANG'
California Arhoretum Foundation, Inc., Arcadia, Calif., and
Los Angeles StSe College, Los Angeles, Calif.
All species of ^Pieris which occur in western North America were
named in the nineteenth century at which time wing coloration and
pattern were the most obvious characteristics used for identification.
Great variations are present between various species of the genus Pieris
with respect to these characters. Such variations have contributed to
disagreement concerning the number of species which occur in North
America. Some authors list seven species: P. rapae, P. napi, P. vir-
girdensis, P. sisymhrii, P. beckeri, P. pmlodke and F. occidentdis ( Klots
1932, McDunnough 1939) . Abbott et ai ( I960) in a statistical study
utilized six morphological characters which included the length and
the width of the front wing, the length of the hind wing, body length
and the degrees of melanization of the upper and lower sides of the
wings of the specimens. They found that there is a continuous varia-
tion in wing sizes, color and marking between P. protodice, P, occi-
dentdis and P. sisymbrii. Such variations were found associated with
geographical location and climate. They suggested that F. protodke,
P, occidemtdis and F. sisymbrii were a single species. Klots (1961) in
Hs ktest Pieris list, cited only six species, excluding F. occidentdis.
Hovanitz (1962) combined the distribution of P. occidentdis • mA
P, pfotodice in a geographical study because of uncertainty of how to
separate them. Evidence for considering them as separate species is
given in this paper. Hovanitz (1963) also indicated that F. mpi and
P. vifgmiensis ought to be considered as a single species at the present
time for nomenciatiiral purposes pending more complete evidence of
their possible sympatric distribution. Since F. vwgimemis occurs only
in the region east of the Rocky Mountains in North America, it is
excluded from consideration in this paper. In the present paper, mainly
morphological characters are studied, m^hich are intended to aid in
the elucidation of the biological relationships between these various
species of Pieris.
^This work was supplementary to work aided by a grant from the National
Science Foupdation.
97
98
CHANG
]. Kes. Lepid,
MATERIALS AND METHODS
Five hundred specimens have been studied from the collections
gathered by William Hovanitz and the author throughout the region
in recent years. Despite this fact, this study is not intended of an
exhaustive geographical study.
The wing characters used in this study include: (1) wing color
and pattern^ of which the detailed description will not be given; how-
ever, the most important characters are outlined in the text, and
(2) wing venation, which is relatively uniform in the genus Pieris.
Differences have been found in the length of the Ry, and in the location
of R2 in respect of M2 and (3) the androconial scales which are the
scent. scales found with the black scales in the dot on or near the discal
vein on the male fore wings.
The characters used for the male genitalia include ( 1 ) the uncus,
(2) the juxta, (3) the saccus and (4) the penis. For the fem?le
genitalia, only the signum bursa has been used; this is a chitinized
structure on the corpus bursa.
The overall size of the insects influences the size of their internal
structures. To make this size difference of negligible influence, the
simple ratio: the length of the left front wing against the length of a
particular structure has been used for aiding in the elimination of
such possible differences.
Ratio
the length of a given structure ^ ^00
the length of left front wing
The terminology of structural parts employed here has been adapted
from Klofs (1931). The color terms whenever used in the text follow
the "Dictionary of Color” by Maerz and Paul (1930).
L P. rapae and P. napi
Color and pattern (fig. la): P. rapae and P.-napi are different
from other species by lacking a dot on the discal vein of the fore wings.
Most samples of P. rapae and P. napi are easily separated, except the
all-white forms of both species.
Venation: The mean ratio (table 1) between the length of the
left fore wing and the length of the discal cell are statistically differ-
ent for these two species. The probability that the difference might
be due to chance alone is less than 0.001 when the degree of freedom
is 45 ( as tested by ) . The histogram ( fig. 3 ) shows however that
the range of ratios is continuous from one species to another. In
actual fact, however, 00 overlap has been observed in the investigation
between these two species though it may occur when the sample size
is larger.
Location of the discal cell, and method of measuring venation are
illustrated in fig. 2.
2(2):97-l25, 196}
MORPHOLOGY OF PIERIiS
99
Fig. la.
Pieris rapae and Pieris napi. Male on left, female on right. Upper
four are upper side;lower four are lower side.
100
CHANG
]. Res. Lepid.
Fig. lb. Pieris beckeri and Pieris sisymhrii. Male on left, female on right.
Upper four are upper side, lower four are lower side.
2(2):97.12i, 1963
MORPHOLOGY OF PIERIS
101
Fig. Ic. Pieris protodice and Pieris occidentalis. Male on left, female on right.
Upper four are upper side, lower four are lower side.
102
CHANG
]. Res. Lepid.
Se D, = Standard deviation; S, E. = Standard error
2(2):97-125, 1963
MORPHOLOGY OF PIERIS
103
Androconia (fig. 4, 5, table 2): Both P. rapae and P. napi have
androconial scales with broadened tops and two lobes. A difference is
found on the width of the scale’s stalk which for P. rapae is narrower
than P. napi. The probability as shown by the 'P” test that the differ-
ence between the mean ratio of these two species is due to chance
alone is less than 0.001 for a degree of freedom of 40. The histogram
(fig, 5) shows that there is a wide gap between the ratios. This is one
of the characters that can be used to separate the all white forms of a
P, rapae and P. napi.
Male genitalia: (1) the uncus (figs. 6, 7, table 3) is more slender
and less curved at the ventral side for P. rapae and P. napi than for the
other four species in the genus Pieris. The ratios: the length of the
uncus against the length of the fore wing range from 2.07 to 2.84 for
P. rapae; and frorn 2.69 to 3.05 for P. napi. The probability that
this difference might be due to chance alone is 0.025 for a degrees
of freedom of 18 as tested by t:
Fig. 2. Location of discal cell and points of measurement
DISCAL CELL
104
CHANG
I. Ii,s. LcpiJ.
j
O
Q
(a
kj
k
Ui
>J
1
a:
-o
g
lo
(%J NOUVindOd
Pier is eapf-'
'Pier is sisymbrfi
Fig, 4. Androconial scales (400x)
ANDROCONIA
106
CHANG
]. Res. Lepid.
(%) NOlivnndOd
o
o
X
istogram illustrating the differences in ratios distribution between
, rapae and P. napi
107
2(2):97.12K 196} MORPHOLOGY OF PIERIS
m
d
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4J
m
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108
CHANG
J. Kes. Leptd.
(2) The juxta (fig. 7) is quite different in shape between P.
rapae and P. napi. The lower end of the juxta of P. rapae is triangular;
however, in P. napi, this structure is reduced in size and not hollowed
out.
(3) The saccus and penis (fig 8, table 4) show no significant
differences between P. rapae and P. napi,
Signum bursa ( fig. 8 ) : This structure is again a good character
to separate P, rapae and P, napi. The signum bursa of P. rapae is like
a pair of kidneys; the central part is less toothed. In P, napi, it has
a long unchitinized tail which is so distinctive that there is no problem
in separating this species from others in the genus Pieris.
The samples of P. rapae came from ( 1 ) Arcadia, Los Angeles Coun-
ty, California (5 individuals), (2) Newport Beach, Orange Co., Calif.
(5), (3) Riverside, Riverside Co. Calif. (5), (4) Bishop, Inyo Co.
Calif. (5), (3) Klamath Falls, Klamath Co. Oregon (5), and (6)
Satus Creek, Yakima Co. Washington ( 5 ) . The samples of P. napi
came from (1) Lopez Canyon, San Luis Obispo Co. Calif. (5), (2)
Berkeley, Calif. (1), (3) Utah (1), (4) Flurricane Ridge, Olympic
National Park, Wash. (10), (6) British Columbia (2), (7) Yukon
Territory (6), and (8) Alaska (3).
Fig. 6. The lateral view of male genitalia
2(2):97-i25, 196) MORPHOLOGY OF PIERIS
Fig. 7. Male genital structures
109
Table 4 Ratios between length of various genital structures and length of front wing
110
CHANG
/. Res. Lepiil.
Probability
+
o
+
«
o
o
o
o*
■4^
00
cn
•
O'-
«
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(M
W
m
0, 09
0. 07
60 *0
01 *0
0* 04
0. 04
Q
m
fM ^
• •
o o
0. 28
0, 24
0. 31
0. 19
Mean of
ratio
rj o
• m
O t-”
« •
00 m
» «
ra fP)
Mean length
of front wing
g
g
O'
m 00
V
tM
00
r-
O* ro*
22.96
23. 54
Mean length
of saccus
0, 51 mm
0. 51
« — )
• •
O O
00
• #
o o
o
a- r-
00 00
53
30
Species
rapae
napi
sisymbrii
beckeri
protodice
occidentalis
a
ro
O
Probability
O
o
o*
o
o
•
o
cn
00
•
#
4^
r«-
r-
W
fM
PO r~4
m
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m
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m
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pH
o
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pH s-H
m *
9 #
bJO
d
g g
s g
0
M H
f=^
M N
f=s H
fH
6 g
g g
a
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d
0 fO
00 fM
o* o*
O* O*
rd
4^
m bo
fl d
R G
s s
0 ^
6 B
G 6
N N
G K
PO Pj
fO O'
r- 00
r~
0 o
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rM ra
fM fM
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f%J p>4
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P-t .-1
pM fO
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^ 0
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rts
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0.*
d d
2(2)^97-12K ^963
MORPHOLOGY OF PIERIS
111
P E N i S
S i 6 MU M
b’ U R S A
pier is sisymtrii
iic e..
Fig. 8. Male and female genital structures
112
CHANG
J. Res. LcpiJ.
II. P. sisymbrii and P. beckeri
1. Color and pattern (fig. lb); These two species are easily
separated by the color and the pattern on the wings. P. beckeri has
the intensified moss green color and P. sisymbrii has the biskra data
color along the vein on the underside of the wings.
2. Venation (fig. 9, table 5): The length of between P.
sisymbrii and P. beckeri differs. The ratio of fore wing length over
R3 for P. sisymbrii ranges from 2.34 to 5.88; for P. beckeri ranges
from 0.95 to 2.62. The probability that the mean for the two popu-
lations could have been drawn by chance from the same population
is less than 0.001 when degrees of freedom is 24 as tested by t.
3. Androconia (fig. 4): The scent scales of P, sisymbrii and
P, beckeri are similar but unlike P. rapae and P. napi. The tiny, lobe-
less scales of P, sisymbrii are nearly parallel at the lateral border, and
the roots are limited to the base. The scales of P. beckeri are usually
irregular shaped and the roots are often occupied half the length of
the scale.
4. Uncus (fig. 7, table 3) ; The uncus of P. beckeri and P. nsymbrii
have hairy processes at the ventral side, particularly in P. sisymbrii.
The actual length of the uncus in the two is much the same, but the
ratio is different (probability 0.005; d.f. 20) due to the constant
smaller size of wing length in P, sisymbrii.
5. Juxta (fig. 7): The juxta of P. beckeri and P. sisymbrii are
similar except the apex is slightly curved upward in P. beckeri.
6. Saccus (fig. 8, table 4): There is little difference in shape or
size with respect to this structure.
7. Penis ( fig. 8 ) : The shape of the penis in P. sisymbrii and
P. beckeri is like the penis of P. rapae and P. napi. However, it is
distinctly different from the penis of P. protodice and P. occidentalis.
The basal protuberance of the penis of P, beckeri and P. sisymbrii is
insignificant. But it rises abruptly for P. protodice and P. occidentalis.
8. Signum bursa ( fig. 8, table 6 ) : The signum bursa of P. sisym-
brii and P. beckeri is stick-shaped, with large, heavily chitinized teeth.
The difference between the ratios is probably caused by chance error
due to the extremely small sample of P, sisymbrii examined, rather than
to a real difference.
The samples of P. sisymbrii came from ( 1 ) Kelso Valley, Kern
Co. Calif. (2), (2) Roads End, Tulare Co. Calif. (1), (3) Kings
Canyon, Fresno Co. Calif. (2), (4) Hurricane Ridge, Olympic Na-
tional Park, Wash. (6), and (3) Whitehorse, Yukon Territory (1).
The samples of P. beckeri came from ( 1 ) Long Valley, Mono Co.
Calif. (1), (2) Mono Lake, Mono Co. Calif. (2), (3) Gardnerville,
Douglas Co. Nevada (2), (4) Minden, Douglas Co. Nevada (2),
(3) Doyle, Lassen Co., Calif. (2), and (6) Sams Creek, Yakima Co.
Wash. (3).
Table 6 Ratios between length of various genital structures and length of front wing
2(2):97-I2'>, l')6)
MORPHOLOGY OF PIERIS
IH
d
bO
*r4
m
Probability
0
•
0
0
0
•
0
sO
00
•
•
CO
I— <
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ra r-
N
CO irt
CO m
0 0
0 0
•
• •
• •
CO
0 0
0 0
m
Q
m
0 m
0
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•
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0 0
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fH 0
0)
• •
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N
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00 00
CO 0
§ 5
• •
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(S3 (S3
l-s
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at
Pi
^ J
4J
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be
d
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g
0 p
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• •
• •
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0 0
0 0
Vi
0
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10 m
Z
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i ^
0
4-> -fI
4)
m 0
0 t»
P.
* 9)
P( 0
CO
m ^
p 0
Q
rd
m
PI
0)
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d
O
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rt
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W
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CO
pcj
•rH 'd
d p
o
d)
P4
CO
0
o
o
o
Pi
p.
114
CHANG
]. Res. Lepicl.
III. P. protodice and P. occidentalis
1. Color and pattern (fig. Ic, table 7): The color and pattern
of these two species are many times undistinguishable, especially in
comparison between the high temperature form of P. occidentalis and
the low temperature form of P. protodice. The primary difference in
the pattern, besides the gross one of more extensive black pigment on
P. occidentalis males than on P. protodice males, is in the size and
shape of the dot in the tornus area near the inner margin on the
underside of the fore wings. The dot on P. protodice is large, square
shaped and not clearly outlined, part of the dot being sometimes sub-
merged on the uppersides. The dot of P, occidentalis is small, relatively
clearly outlined, usually not submerged. The ratios for the width of
the tornus dot range from 8.73 to 15.38 for P. protodice; and from 3-95
to 8.51 for P. occidentalis. The probability that this difference is due
to chance alone is less than 0.001 for the degrees of freedom of 60
by the t test.
2. Venation (fig 9, 10, table 5); The R:^ veins of P protodice
and P, occidentalis are quite short, sometimes being completely miss-
ing. The mean ratios of wing length over length between the two
species do not show much difference. (1.17 for P. protodice\ and 1.15
for P. occidentalis). However, the difference is significant between
P, protodice and P. sisymhrii in comparing the ratios despite ^the sim-
ilarities in their wing color and pattern. The mean ratio for P. sismyhrii
is 4.16; and for P. protodice it is 1.17. That the difference as indicated
might be due to chance is less than 1/1000 for the degrees of freedom
of 46 by the t test.
In P, protodice, the intersection of Rl>, Rs+Mi (point "A”) is usual-
ly above point "B” where M2 joins that vein; hov/ever, point "A” is
usually on or below point "B” in P. occidentalis. The distance between
point "A” and point "B” is measured and plotted on the diagram
(fig. 11) against the ratio obtained from the width of the tornus dot
to the length of the fore wing. When point "A” is above point "B”
the measurement is given a "-b” sign. When point "A” is below point
”B” the measurement is given a '' — ” sign. The diagram (fig. 11)
shows that P. protodice and P, occidentalis are well separated by these
two characters. Few exceptional dots on the diagram as the measure-
ments between point "A” and point "B” having a ” — ” sign for P.
protodice or vice versa for P. occidentalis are from the specimens col-
lected at the overlap zone of these two species.
The width of tornus dot and the location of R2 in respect of
M2 are two pirmary characters used in the investigation to separate the
two species.
3. Androconia: Thirty-five specimens of male P. protodice and
twenty-eight specimens of male P. occidentalis from ten different
localities were checked, but no androconial scales have ever been found.
Similar results were also observed by Bernard! ( 1947) .
2(2);97-125, 1963
MORPHOLOGY OF PIERIS
115
P . protodice
Fig, 9. Venations showing the differences in the length of R3
116
CHANG
]. Res. Lepid.
prim ary
P. protodice
Fig. 10. Venations showing the left fore wing and the differences in the
location of point "A” in respect to point "B” between P. protodice
and F. Occident alts
2(2):97-l2^, 196}
MORPHOLOGY OF FIERI S
117
4. Male genitalia: (1) Uncus and juxta (fig. 7, table 3): No
difference is found between P, protodice and P, occidentalis in respect
to these two characters.
(2) Saccus (fig. 8, 12, table 4): The length of saccus is statis-
tically different between P. protodice and P. occidentalis. The mean
ratio (length of fore wing over length of saccus) for P. protodice is
2.68; for P. occidentalis it is 3.45. The probability is less than 0.001
that the difference may be due to chance when the degrees of freedom
is 83 as shown by the t test.
The .scatter diagram (fig. 12), which shows each individual ratio
of these two species against its own wing length, shows that overlap
is present in this character despite the difference in mean ratio. This
may be due to the ‘variation in wing size caused by geographical and
climatical difference. In cooler areas, the butterfly is usually smaller
and in warmer areas, larger. If some specimens of P. occidentalis are
from the extreme southern locations and some specimens of P. protodice
from the extreme north, the gap in ratio between these two populations
is reduced, and overlap occurs. This point is demonstrated by fig. 14.
If the specimens of P. protodice and P, occidentalis are from the same
locality and collected at the same time, these two species are usually
separable.
( 3 ) Penis ( fig. 8 ) : The penis of P. protodice and P, occidentalis
is strongly curved at the basal part and this difference makes these two
species easily distinguishable from other species in the genus Pieris.
The penis size in P. protodice is smaller than in P. occidentalis.
5. Signum bursa (fig. 8, table 6): The signum bursa of P, pro-
todice and P, occidentalis is similar to P, sisymbrii, but with smaller
teeth and less chitinization. The length of signum bursa in P. protodice
is shorter than the one in P, occidentalis': The mean ratio (length of
fore wdng over length of signum bursa) for P. protodice is 3.18 and
for P. occidentalis is 4.03. The probability that this difference is due
to chance alone is less than 0.001 when the degrees of freedom is 60
by the t test. The scatter diagram (fig. 13) for individual ratios shows
that there is no overlap present between these two species with respect
to this character.
The mean ratio of each population at a given locality for P. proto-
dice and P. occidentalis is also plotted on a diagram (fig. 14) which
shows that the ratios of the specimens from northern localities such
as Alaska are larger than the ratios of the specimens from southern
localities. The differences in ratios for the specimens from various
southern localities are not significant. This is porbably due to the
closeness in distance between the localities and to similarities in en-
vironment. This diagram illustrates however that the ratio is not every-
where unchanged. The ratio gradually increases in the more northern
VENATION
P. PROTODICE (k)
P. OCCIDENTALIS { •)
118
CHANG
/. Res. Lcpid.
001 X
iOO SnNMOi HiOIM
Fig. 11. The measurements between point "A” and point "B” (see fig. 10)
plotted against the ratios obtained from the width of the tornus dot
to the length of the fore wing
S AC C U S
2(2):97-125, 196}
MORPHOLOGY OF PIERIS
119
»(+++
-++ + + +
44
4- 4f 4-4-
Q. (x:
MJ JO H 19 N 3 1
00! X
snooir s jo h i9N3 i
Fig. 12. Scatter diagram showing the individual ratio of saccus length to the
length of fore wing plotted against the individual saccus length in
P. protodice and P. occidentalis
V s y ns
120
CHANG
/. Ke%. Lepid.
o
Q
^ k
k,
Q
SS
!>>
©
<0
o
to
o
©
00 ! X ^ ^ NI9N3 1
8 § J O HI 9N31
Fig. 13. Scatter diagram showing the individual ratio of the length of signum
bursa to the length of fore wing plotted against the individual
length of signum bursa in P. protodice and P. occidentalis
2 (2) -.97 -125, 1963
MORPHOLOGY OF PIERIS 12 i
■M'J HI3 1 N¥3m
MIS M3 7 NV3M
MEAM LENGTH OF SIGNUM BURSA (mm)
Fig. 14 Scatter diagram showing the mean ratio of the length of signum
bursa to the length of fore wing in each- locality plotted against the
mean length of signum bursa
122
CHANG
J. Res. Lepid.
localities. The ratios of two species may overlap at different localities
though they do not do so at the same location.
The samples of P. protodice came from ( I ) Laguna Canyon, Orange
Co. Calif. (11), (2) Moorpark, Ventura Co. Calif. (4), (3) Inde-
pendence, Inyo Co. Calif. (2), (4) Big Pine, Inyo Co. Calif. (2),
( 3 ) Mammoth Lake, Mono Co. Calif. ( 1 ) , (6) Mono Lake, Mono Co.
Calif. ( 4 ) , (7) Coleville, Mono Co. Calif. (5) , (8) Gardnerville
Douglas Co. Nevada (1), (9) Minden, Douglas Co. Nevada (1),
and (10) Doyle, Lassen Co. Calif. (4). The samples of P. occidentalis
came from (1) Long Valley, Mono Co. Calif. (1), (2) Mammoth
Creek, Mono Co. Calif. ( 2 ) , ( 3 ) Mono Lake, Mono Co. Calif. ( 2 ) ,
(4) Coleville, Mono Co. Calif. (1), (3) Gardnerville, Douglas Co.
Nevada (11), ( 6 ) Doyle, Lassen Co. Calif. ( 3 ) , ( 7 ) Klamath Falls,
Klamath Co. Oregon (4), (8) Satus Creek, Yakima Co. Washington
( 2 ) , ( 9 ) British Columbia ( 1 ) , and (10) Alaska ( 1 ) .
GEOGRAPHICAL DISTRIBUTION OF P. PROTODICE
AND P. OCCIDENTALIS
The geographical distribution of all species in the genus Pieris in
North America except P. protodice and P, occidentalis is thoroughly in-
vestigated by Hovanitz (1962) (fig. 15). The data here presented are
mostly gained by extensive search throughout the range of P. protodice
and P. occidentalis in West Coast conducted by William Hovanitz in the
summer of 1962. P, protodice is most abundant in the south, from
the southern tip of Baja California northwards, gradually reducing its
population and being replaced by P. occidentalis. The northern border
of P. protodice is still unclear. At present this species is only in Cali-
fornia and Nevada southwards. There is no record in Oregon and north-
ward. P, occidentalis extends from central Alaska along the Rocky
Mountains and the Cascade Mountains southwards into the Sierra
Nevada Mountains (and the Rocky Mountains). The most southern
locality known from our data at present in the Pacific coast area is
Long Valley, Mono Co., Calif. It is very likely this species may be
found at more southern places on high Sierra Nevada Mountains.
P, occidentalis and P. protodice overlap from Long Valley, Mono
Co. Calif, northward to Doyle, Lassen Co., Calif. Further investiga-
tion may extend the overlap zone more northward even to Washington.
The habitats of these two species are very similar; both like sunny,
open, grassland. P, protodice prefers warmer temperatures and semi-
desert conditions. It feeds on Brassica nigra^ Caulanthus sp., Lepidium
densiflorum, Sisymbrium altissimum, Cleome lutea, and T hely podium
lasiophyllum. P. occidentalis prefers relatively cool temperatures. It
is usally found at places relatively moist, as near a lake, ditch or farm
land. The food plants of P. occidentalis are nearly the same as for
2(2):97-l2S, 196}
MORPHOLOGY OF PIERIS
Fig. 15.
Map showing the distribution of P. protodice and P. occidentalis
western North America
124
CHANG
). Res. Lct)iJ.
P. protodice except that Cleome Intea has not yet been recorded.
P. protodice and P. occidentalis are found flying together in the
overlap zone. The males of these two species at a particular locality
are usually separable by the marking and color on the wings, in which
P. protodice is less darkly marked than P. occidentalis.
DISCUSSION AND CONCLUSIONS
The present data indicates some clear structural differences be-
tween the six species of Pieris. P. rapae and P. napi are similar in
many characters, such as uncus, saccus, and the color and pattern on
the wings, but they also can be distinguished by their differences in
pattern and the signum bursa, juxta and androconial scales. P. sisymbrii
and P. heckeri are similar in their uncus, saccus, androconial scales
and signum bursa, but can usually be distinguished by wing pattern
and venation. P. protodice and P. occidentalis can be distinguished by
wing pattern, wing venation, structural differences in the length of
saccus and signum bursa only in a statistical study. These two species
also differ in their geographical distribution pattern, but overlap greatly
in a sympatric area.
P. sisymbrii differs in many characters from P. protodice, such as
wing venation, genital structures and others. The most significant
difference is the presence of androconial scales, v/hich are absent in
P. protodice and P. occidentalis.
The results serve to show that P. protodice and P. occidentalis are
sympatric species, an uncertainty which led Hovanitz (1962) to com-
bine their geographical distributions into one map. The other species
of the genus Pieris in Western North America are P. rapae, P. napi.
P. sisymbrii, and P. beckeri. It is always possible that even these may
be found at some time in the future to be subdivided into cryptic
species by some characters not yet discovered.
ACKNOWLEDGEMENT
The author wishes to express his deep appreciation to Dr. William
Hovanitz and Dr. E. 1. Schlinger for their help and constant advice
during the entire study.
2(2):97-l2‘>, 196}
MORPHOLOGY OF PIERiS
125
LITERATURE CIlTD
ASAKAWA, F. 1961. Morphological observation on the androconia in the
butterflies. (Ja) New Ent. 10(2): 7-15.
ABBOTT, W., L. S. DILLON, and R. R. SHRODE. I960. Geographic
variation in Pieris protodice Boisduval and Leconte ( Lepidoptera : Pieri-
dae). Wasmann J. Biology ^ 18: 103-127.
BERNARDI, G. 1947. Revision de la classification des especes holarctiques
des genus Pieris Schr. et Pontia Fabr. (Pep., Pieridae). Miscellanea En-
tomologica, Paris 44: 65-80.
HOVANITZ, W. 1962. The distribution of the species of the genus Pieris
in North America. /, Res. Lepid. 1 (1): 78-83.
• . 1963. The relation of Pieris virginiensis Edw. to Pieris napi
L. Species formation in Pieris.^ J. Res. Lepid. 1 (2): 124-134.
JACKSON, R. A. 1946. Causes for seasonal variation in the number of
Lepidoptera. Proc. & Trans. S. London Ent. & Nat. Hist. Soc. 1945-46:
43-51.
KLOTS, A. B. 1931. A generic revision of the Pieridae (Lepidoptera) to-
gether with a study of the male genitalia. Entomologica Americama 7
(new series) (3): 139-204; (4): 205-243.
KLOTS, A. B. 1961. Family Pieridae. In "How to know the butterflies”
edited by Ehrlich, P. R. and A. H. Ehrlich. Wm. C. Brown Co. Du-
. buque, Iowa.
MAERZ, A. and M. R. PAUL. 1930. A dictionary of color. 1st Ed. Mc-
Graw-Hill Book Co. New York.
MARIANI, M. 1937. Anatomia e fisiologia degli organ! genitali femminili
delle Pieris Schrk. (Lepidoptera, Pieridae). Festschrift fur Prof. Dr.
Embrik Strand. 3: 434-450.
McDUNNOUGH, J. 1939- Check list of the Lepidoptera of Canada and
the United States of America. Mem. So. Calif. Acad. Sci. 11(1) : 1-171.
McHENRY, P. 1962. The generic, specific and lower category names of
the Nearctic Butterflies. Pat 1. The genus Pieris. J. Res. Lepid 1(1):
63-71.
OKANO, M. 1951. Comparative morphology of the male genitalia of
Japanese Pieridae. Ann. Report Gakougei Eac. livate Univ. 2: 38-46.
PETERSON, B. 1963. The genitalia of some colias species. /. Res. Lepid.
1 (2) : 135-156.
QUERCI, O. 1936. Notes on Pontia protodice Boisduval and Leconte (Lep:
Pieridae). Trans. Amer. Ent. Soc. 62: 37-47.
TUXEN, S. L. (editor). 1956. Taxonomist’s glossary of genitalia in insects.
Ejnar Munksgaard, Copenhagen.
WARREN, B. C. S. 1961. The androconial scales and their bearing on the
question of speciation in the genus Pieris, (Lepidoptera.) Ent. Tidskr.
82(3/4) : 121T48.
126
CKANG
). Res. Lepid.
BIOGRAPHICAL SKETCHES
CHANG, VINCENT CHUEN SUN
[2855 Nina St, , No, 2, Pasadena, California]
Born: Shanghai, China, March 17, 1935
B„ A, : The Agriciiltural Institute of Nanking, Nanking, China,
1957
Graduate student: University of California, Riverside and Los
Angeles State College
Position: Research Assistant, California Arboretum Foundation,
Inc,, 1959-63
Interests: All aspects of Pieris,
including ecology, phy-
siology, and morphology
Photography of activities of
butterflies
Publications: A series of papers on
food preferences of Pieris
Journal of Research on the Lepidoptera
2(2) :127-136, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 196}
GENETIC AND ENVIRONMENTAL VARIATION
IN PIERIS BRASSICAE
BRIAN O. C. GARDINER
Agricultural Research Council, Unit of Insect Physiology,
34A, Storey’s Way, Cambridge, England
INTRODUCTION
The principal form of variation in Pieris hrassicae L. consists
of either a reduction or expansion in the amount of black present. In
certain local populations the variation is constant and marked enough
to be designated as a race, such as P. hrassicae cheiranthi in the Canary
Islands. Marked variation in color, however, is exceedingly rare and,
until now, only some half dozen specimens were known.
In 1950 a stock of P. hrassicae originating from Cambridge was
established at the Unit of Insect Physiology and has been continuously
bred ever since. This stock has already been reported on (David &
Gardiner 1951; David 1957). In 1959 another stock, originating from
Oxford, was established as a separate culture and was also continuously
brooded under the same conditions as the Cambridge stock.
Until i960 the amount of variation noticed in the Cambridge
stock was simall, and confined to known variation in the black markings.
This has already been briefly reported on (David «& Gardiner 1961).
During the past three years, however, two color varieties have ap-
peared, as well as the previously unknown albino form; known varieties
also seem to have been more common, although this could well be
due to more careful observation (fig. 1).
The opportunity was taken to determine the genetics of these
varieties and also to examine the combination of two pairs of allelo-
morphs to produce a double recessive. It is believed that this is the first
time such a cross has been done with a butterfly.
From time to time specimens considerably smaller than the normal
size turn up in various species of butterflies. In the case of P. hrassicae
specimens of between 40-50 mmi. wing span are referable to ab. minor
Ksienschopolsky. According to Frohawk (1934) normal wild caught
specimens vary from 63 mm. in the male to 76 mim. in the female.
In the continuously brooded culture kept at the Unit of Insect Physi-
ology the size is 55-63 mm. in the male and 54-65 mm. in the female
(David & Gardiner 1961).
127
128
GARDINER
/. Res. Lepid:
Occasional specimens of ab. minor were noted in the Cambridge
stock. At times entire batches have been on the small side. Specimens
bred from these were normal. It occurred to the author that ab. minor
might be an effect of starvation rather than a genetic effect. This
has been investigated as will be described.
DESCRIPTIVE
Pieris hrassicae L. ab. coerulea ab. nov.
The normal cream and green coloration on the underside of the
wings is replaced by a pale blue color. On the upper side, the white
areas of the wings have a pure white appearance and are rather thinly
scaled; in normal specimens they are creamy white or off white in color.
Larvae and pupae normal. The majority of the adults of this variety
fail to eclose properly. Some specimens may emerge with striped
scales, especially on the hind wing, which gives them a translucent'
blue color, deeper in shade than normally formed coerulea.
Holotype ^ , Allotype $ : Two specimens selected from the F i
generation obtained by the pairing of two ab. coerulea which emerged
from a continuous brooded culture kept in the Laboratory since 1950.
In the author’s collection.
Paratypes: Five pairs also selected from the Fi. Two pairs have
been deposited in the Rothschild-Cockayne-Kettlewell collection at
Tring, two pairs are in the author’s colletion, and one pair in the coL
lection of Mr. H. Douglas Bessemer.
Type locality: The original stock from which this aberration arose
was collected in the vicinity of Cambridge, England, in 1950.
Synonyms, ab pallida. Graham-Smith & Graham-Smith
ab. anthrax. Graham -Smith & Graham-Smith
Graham-Smith and Graham-Smith (19.30) after describing an-
thrax state "Perhaps a specimen exhibited by Leeds (1909) with
undersides of the hind wings a very distinct blue . . . are examples
of this aberration.’’
Under pallida they state "A very marked example from Monk’s
Wood, Hunts., is figured by Frohawk ( 1914, pi. 3, fig. 20)
The specimen figured by Frohawk (1914) is stated by him to have
been taken by Mr. H. A. Leeds in Monk’s Wood in 1906 and there
can be little doubt that it was this specimen that was exhibited by
Leeds at the Annual Exhibition of the South London Entomological
and Natural History Society in 1909-
Mr. A. L. Goodson has compared the present variety with the an-
thrax and pallida in the R.-C.-K. collection at Tring and it in no way
resembles either of them nor, in the author’s opinion, does it bear any
resemblance to the illustrations of anthrax and pallida in the paper by
Graham-Smith and Graham-Smith (1930).
2(2):127-IH, 1963
PIERIS BRASSICAE VARIATION
129
Mr. H. Douglas Bessemer, who has the original Leeds specimen in
his collection, very kindly invited the author to visit him and examine
it. There is no doubt that it is the same as the present variety and was
wrongly' identified by Graham=Smith and Graham-Smith.
Pieris brassicae L. ab. jmmi ab. nov.
In May 1961 it was noticed that the underside of some butterflies
being reared in the Laboratory was different in color to normal speci-
mens, being a pale straw color compared to the normal greenish-yellow.
The difference is more striking in freshly emerged living specimens
than in dead set ones. The larvae are normal but the prepupae and
pupae are a golden-yellow without any trace of the normal green. The
pupae are also without the normal black speckling. Unlike some other
color varieties the butterflies edose normally.
Holotype , Allotype ’$ : Two specimens selected from the culture
in May 1961. In the. author’s collection.
Paratypes: Two pairs similarly selected. In the author’s collection.
Type locality: The original stock from which this aberration arose
was collected at Oxford, England, in August 1959.
Pieris brassicae ab. alhinensis Gardiner
This variety has already been described (Gardiner 1962). It is a
simple recessive and is noted here since it was used with coemlea to
obtain the double recessive.
EXPERIMENTAL
Most insects were bred according to methods already described
(David & Gardiner 1952; David 1957) with the exception that when
a single pair of butterflies was being paired a small nylon covered
cage measuring 1 x 1 x lid feet was used instead of a large cage. A
percentage of the larvae were also reared in 2 lb. jam jars at the rate
of about 20 per jar. This enabled an accurate check to be kept on
mortality. This was generally done with the F| broods, the larger
numbers of the F2 being reared in cages. No difficulties were encount-
ered with either pairing or oviposition nor, apart from the Fi of ab.
jauni, was there any appreciable mortality of the larvae. Due to limit-
ations of time, space and food, it was only possible to rear a small
percentage of the total ova obtained. Since a single fertile brassicae
was found to lay over 500 ova, the potential F2 was some 125,000 in-
dividuals, an impossible number to rear.
ab. coerulea
In November 1962 a female emerged in the Cambridge stock. She
was paired to a normal male from the same batch and the resulting
ova were reared to give an Fi of normal insects. These were crossed
and a selection of the resultant ova reared.
130
GARDINER
]. Kcs. Lepiil.
The F2 produced 347 normal and II6 coerulea butterflies, a ratio
of 2.99 : 1, a very good agreement with the expected 3 : 1 ratio for
a simple recessive. The majority of the coerulea however failed to
eclose normally and remained stuck by the wing tips to the pupal case.
While this brood was being reared, further coerulea emerged from
a batch of the stock Cambridge butterflies. Two pairings v/ere obtained
and all the Fi and subsequent generations were coerulea. This aberration
is therefore a recessive. It’s extreme rarity under natural conditions
can be explained by the difficulty the butterflies have in getting free
of the pupal case.
The blue color is due to an absence of the normal yellow pigments
and is present normally in the wing-membrane. This can be shown
faintly by rubbing off the scales of a normal hrassicae between finger
and thumb. A very similar appearance to coerulea can also be pro-
duced by immersing a normal hrassicae in the bleaching agent sold,
under the name of 'Parazone.’ The question of the chemistry of this
and other color forms of hrassicae is at present under investigation
by a colleague of the present author and will be dealt with in a later
paper.
The failure to eclose is more marked in the males than the females.
In the first two generations only about two per cent of the specimens
could be described as immaculate. About 50 per cent fail to clear the
pupal case and are stuck by one or both fore wing tips; others get
clear but one or more wings fail to expand. In many instances there
is extensive stripping of the scales, especially of the hind wing, which
then has a deep blue translucent appearance.
After several generations the percentage of normal eclosions rises
to about 80 per cent. This is similar to the trend that occurred in
alhinensis (Gardiner 1962). Since 1961 alhinensis has been passed
through a further ten or twelve generations and the percentage failing
to eclose normally is now negligible, and there seems little doubt that
coeridea will eventually reach this stage.
While failure to eclose is a semi-lethal factor linked, initially,
with coerulea it is also partially controlled by the temperature at which
eclosion takes place. Two experiments were done on this, using the Fo
generation of the pure line in the first and alhinensis in the second.
A cage of coerulea pupae was kept at 20 °C. until 12 normal
(5.7%) and 198 crippled specimens had emerged. The remaining
pupae were than transferred to 12.5°C. and at this temperature 38
normal (27.0%) and 103 crippled specimens emerged.
In the second experiment alhinensis was used, half the pupae being
at 25°C. and half at 12.5°. At 25°C. 15 normal (30.0%) and 35
crippled emerged while at 12.5°C. 28 normal (58.4%) and 20 crip-
pled emerged.
These results clearly show that failure to eclose is more marked
at higher temperatures.
2(2):127-1U, /96.!
PIERIS BRASSICAE VARIATION
131
ab jaunt
These were noted in May 1961 in a stock obtained from Oxford
which had been bred for a number of generations in the Laboratory.
A pair of these butterflies was mated and 100 of the ova reared.
In this generation the mortality, mainly granulosis virus, was 76%.
The 14 adults obtained were all of the variety. These were paired at
random and 100 Fo ova reared to produce 97 butterflies, also of the
variety.
After the Fi males had paired with their sisters they were paired
with normal females and a percentage of the ova laid were reared to
produce all normal butterflies. These were mated at random to produce
the F:> of which a percentage of the total ova laid were reared, with
little mortality. This generation produced 100 normal and 34 ab jauni
butterflies. This is a 3 : 1 ratio.
This variety seems to be due to the lack of the normal greenish
colored pigment. It segregates as a simple recessive and breeds true.
It is of interest as being one of the few known instances in the butter-
flies in which a single gene manifests its effect not only in the adult
but also in the earlier stages.
ab minor
About 200 final instar larvae, which had been kept well supplied
with food during their earlier stages, were allowed to feed normally for
3 days and were then starved. About half died, the remainder wander-
ed around their cage and eventually pupated. Normally, at 20 °C,
the final instar larvae feeds for 5 days before pupation ( David & Gard-
iner, 1962).
The majority of the pupae produced butterflies, all ab minor, the
smallest specimens having a wing span of 37 mm. male, 38 mm.
female.
As a control siblings to the starved larvae were kept and allowed
to feed normally and pupate when ready. These all produced normal
sized butterflies.
Eggs obtained by the pairing of several of these minor were reared
and an adequate supply of food was given throughout their life. Normal
size specimens were produced.
It is evident from this result that ab minor can be produced by a
condition of the external environment, starvation, on the larvae. Never-
theless the size of brassicae is, at least in some instances, genetically
controlled. This will be dealt with more fully in the discussion.
ab nigroviridescens
About a hundred prepupae of the Cambridge stock were put in a
desicator containing water to give 100% R.H. and kept at 20 °C. One
reasonable specimen of nigroviridescens Rocci emerged and two par-
tially crippled adults intermediate between normal and nigroviridescens.
132
GARDINER
]. Res. Lepiil.
No breeding was done from these nor have any similar specimens ever
been noted in the Cambridge stock. It is considered probable, how-
ever, that the nigroviridescens produced was a result of the saturated
humidity of the environment.
ab coerulea/ albinensis
The cross between coerulea and albinensis was performed both
ways. One brood, coerulea $ X albinensis $ being reared by the
author and the brood from the reciprocal cross being reared by Mr.
C. F. Rivers of the Agricultural Research Council’s Virus Research Unit.
In both crosses the butterflies used were from a pure bred line of
the respective recessive.
In both crosses the several hundred individuals of the Fi were all
normal brassicae.
The F^ of both crosses gave a ratio very close to 9 : 3 : 3 : 1 as is
to be expected from the independent assortment of two pairs of allelo-
morphs. The back-cross was also performed to give a 1 : 1 : 1 : 1
ratio. The numbers involved are shown in Table 1.
These results show that linkage, which has not yet been shown to
occur in butterflies (Ford 1946) is not present. Since brassicae has a
relatively large number, 15, of chromosomes (Lorkovic, 1941) this is
perhaps not very surprising. It would have been more interesting to
have crossed coeridea with jauni, since both involve a change in the
same pigment, unlike albinensis which is caused by an absence of black.
Unfortunately by the time coerulea appeared the jauni form had been
lost.
Cross
normal
coerulea
albinensis albinensis
coerulea
coerulea $>
X
albinensis 2
921
295
304
97
albinensis d
X
coerulea $
695
233
227
76
coerulea/ albinensis
X
double heterozygote
97
96
92
95
Table 1. The numbers of butterflies reared in the F2 of the double re-
cessive cross and the Fi of the back-cross.
2(2):127-136, 1963
PIERIS BRASSICAE VARIATION
133
As might be expected the majority of the double recessives failed
to eclose normally but, like their respective grandparents, the percentage
of normal eclosions rises with continual breeding.
Homeosis
This is extremely rare in butterflies. In hrassicae specimens are
known in which the fore wing black markings are duplicated on one
side of the hind wing. According to Ford (1946) there is no definite
evidence of its nature in butterflies but it has been shown to be genetic
in Drosophila.
During the course of hybridising Cambridge stock hrassicae with
race cheiranthi from the Canary Islands, three homeotic specimens
turned up in one brood of some 300 individuals, all the progeny of one
pairing.
The male parent was an F3 from a back-cross of an Fi hrassicae/
cheiranthi hybrid to cheiranthi, while the female was an alhinensis ex-
tracted in the F2 of an alhinensis crossed to the Fi hrassicae/ cheiranthi
hybrid. The cheiranthi used for the back -cross and the Fi hrassicae/
cheiranthi hybrids had the same father.
All the homeotics were male with the black markings on the fore
wing underside partially replicated on the hind wing of one side only.
One male was mated to his sister and sib pairing was continued
to the F;^j some 300 individuals being reared in each generation. No
more homeotic specimens were produced.
This is of course a negative result, but it shows that homeosis is
not caused by a single recessive allelomorph. The most likely explana-
tion of its occurrence, in this instance, is an upset in the genetic com-
patability due to a slight variation between the parent genes.
DISCUSSION
The continuously brooded stock of hrassicae kept at the Unit of Insect
Physiology is now thirteen years old and provides a unique opportun-
ity for the study of variation in a butterfly. The numbers produced
have varied from a fev^ dozen to a thousand or more per week.
In addition to a number of known aberrations two new ones, jauni
and alhinensis, have been produced. A third, coerulea, was only pre-
viously known from two examples, both taken over fifty years ago.
As was to be expected, changes in the color of the wings, coerulea
and jauni, are recessives. It is probable that the yellow form ab flava
Kane is also recessive and its extreme rarity is, like coerulea, due to
failure to eclose normally.
The failure to eclose of coerulea and alhinensis appears to be due
to the wings sticking to the pupal case. Indeed scales can be seen to
be left behind in the pupal cases from which specimens have succeeded
in getting free. It is however a disability which is overcome by con-
Left side, from top down, ab coemlea $ allotype (upperside). $ para-
type (underside L). d holotype (underside). $ coemlea /albinensis (upper-
side) L. Right side. Typical $ ( underside ). ab $ holotype (un-
derside), ab minor $ (upperside). d brassicae/cheiranthi hybrid underside
showing homeosis.
2(2):}27-]36, 1963
PIERIS BRASSICAE VARIATION
135
tinued breeding. This failure to eclose, however, again occurs if albin-
ensis, from a normally emerging stock is crossed to typical hrassicae
and extracted again in the Fi>. The effect is mechanical, more marked
in the males than in the females and is affected by temperature. It
might be that it is a genetic effect and it would be interesting to breed
from the crippled specimens if this were possible. Since only perfect
images are used for breeding, there is a very strong selection towards
normality and only recessive genes producing crippling are rapidly elim-
inated for butterflies which cannot fly are unable to mate or oviposit.
Although minor can be produced by starvation the size of hrassicae
is also genetically controlled. From time to time small backward larvae
are found in stocks of hrassicae. Attempts to breed these have so far
been unsuccessful. Usually the larvae die. Two or three have been
brought to pupation, the pupae being comparable in size to minor, but
no adults emerged.
In 1951 the wing span of the Cambridge stock hrassicae was 53
mm. ^ and 58 mm. $ (David & Gardiner, 1952). By I960 how-
ever this had risen to 58.5 mm. in the male, while the females had
decreased to 57.9 mm. (David & Gardiner, 1961).
The continuous inbreeding of the Cambridge stock has produced
specimens with virtually no size difference between the sexes. This is
smaller, especially in the female than the respective male and female
sizes of 63 mm. and 76 mm. given by Frohawk (1934). It would
appear that Frohawk had some unusually large female specimens before
him. The author now believes that this figure may be a misprint in
Frohawk’s book. A 76 mm. hrassicae would be a giant specimen. Wild
specimens caught in 1943 from the author’s collection being 58 mm.
and 63 mm, while Fi bred examples from a freshly caught wild female
in 1957 are 58 mm. in both sexes.
The Cambridge stock is therefore not significantly different in
size to some wild populations, but both it and wild populations vary
in size over a period of years.
In P. hrassicae race cheiranthi the males are 60-65 mm. and the
females 65-70 mm. (12 examples measured), but the wings are also
broader so that the butterfly looks even larger. If these are crossed to
Cambridge stock hrassicae the Fi insects are interm.ediate in size and
in the F2 a range of sizes are produced. Size in this instance is clearly
under genetic control.
The Cambridge stock hrassicae are on the whole lighter and with
less black than wild populations. The varieties that have turned up,
reducta Fritsch^ colliurensis Gelin and intermediates, are all those with
a reduction of black. ^
The author has in his collection a female series from heavily marked
with a trace of fasciata Kiefer to extreme colliurensis, in which the
remaining spot is very feint.
136
GARDINER
]. Res. Lepid.
The evidence from this and also from hybrid crosses to cbeiranthi
(which will be dealt with fully in a subsequent paper) clearly shows
that a multifactorial effect is involved in the degree of black on the
wings.
SUMMARY
‘ Two new varieties of Pieris brassicae L., coemlea and jauni are
named and shown to be recessive in character.
The failure to eclose in early broods of coemlea and albinensis is
shown to be partly influenced by temperature.
The double recessive coemlea/ albinensis has been reared and the
two allelomorphs concerned shown not to be linked.
Ab. nigroviridescens is considered to be probably produced by
saturated humidity.
The small form, minor, can be produced by starvation. Neverthe-
less size in brassicae is also genetically controlled.
The size of brassicae, both bred and wild, shows variation over a
period of years. It is also believed that Frohawk’s figure of 76 mm.
for the female wing span is a misprint.
Homeosis is shown not to be due to a simple recessive gene.
The Cambridge stock of brassicae shows a tendency for reduction in
the amount of black on the wings. The evidence is that this is multi-
factorial.
REFERENCES
DAVID, W.A.L. 1957. Breeding Pieris brassicae L. and Apanteles glomeratus
L. as experimental insects. U. Pflkrankh. 64: ^12-All.
DAVID, W. A. L., GARDINER, B.O.C. 1952. Laboratory breeding of Pieris
brassicae L. and Apantless glomeratus L. Proc. R. ent. Soc. Lond (A)
27: 54-56.
— 1961. The mating behaviour of Pieris brassicae (L.) in a laboratory
culture. Bull. ent. Res. 52: 256-280.
— — 1962. Observations on the larvae and pupae of Pieris brassicae (L.)
in a laboratory culture. Bull. ent. Res. 53: 417-436.
FORD, E. B. 1946. Butterflies. Collins, London.
FROHAWK, F. W. 1914. A Natural History of the British butterfies.
1934. The complete book of British butterflies. Ward, Locke,
London.
GARDINER, B.O.C. 1962. An albino form of Pieris brassicae (Lep.,
Pieridae). Ent. Gaz. 13: 97-100.
GRAHAM-SMITH, G. S., GRAHAM-SMITH, W. 1930. Pieris brassicae, L.,
with special reference to aberrations from Aberdeenshire. Ent. Rec. &
J. Var. 42: 1-7.
LORKOVIC, Z. 1941. Die Chromosomenzahlen in der Spermatogenese de
Tagf alter. Chromosoma 2: 151T191-
Journal of Research on the Lepidoptera
2(2) :137-141, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 1963
TYPE LOCALITIES OF THE MEGATHYMIDAE
H. A. FREEMAN
1605 Lewis, Garland Texas
It is not possible for a person to make a careful systematic
or ecological study of any group of lepidoptera unless the type local-
ities of all of the known species and subspecies are well known. In
recent years this information has been carefully presented in the orig-
inal descriptions, however some of our earlier authors were not as
careful and we have to do considerable research in order to associate
a present habitat with their findings.
In working with the Megathymidae I have found that there are
several cases where no specific type locality was given or else the
locality was entirely too general. With that in mind this paper pre-
sents a list of these localities (Table 1) with the hope that it will make
it easier for future workers to have the type localities at hand while
they are doing research on this particular group of lepidoptera.
REFERENCES
BARNES, WILLIAM, & JAMES H. McDUNNOUGH, 1912. A review of
the Megathymidae. Contrih. Nat. Hist. Lepid. No. Amer. 1: 1-28, pps.
BASSETT, ANNA S., 1938. Some Georgia records of John Abbot, Naturalist.
Auk 55: 244-254.
BELL, ERNEST L,, 1938a. The Hesperioidea. Bull. Cheyenne Mountain Mus.,
1: 1-35.
• — 1938b. A new genus and five new species of Neotropical Hes-
periidae. Amer. Mus. Novitates, no. 1013: 1-11, figs. 1-6.
BUTLER, ANDREW G., & HERBERT DRUCE, 1872. Descriptions of new
genera and species of lepidoptera from Costa Rica. Cistula Enet. 1 : 95-118.
DOW, ROBERT P., 1914. John Abbot of Georgia. ]ourn. New York Ent.
Soc. 22: 65-72.
Thanks are expressed to the National Science Foundation for research
grants making my work on the Megathymidae possible.
137
138
FREEMAN
J. Res. LcpiJ.
EDWARDS, W. H., 1882. Description of species of butterflies taken in
Arizona by Jacob Doll, 1881. Papilio 2: 19-29.
FREEMAN, H. A., 1943. Notes on and redescriptions of Megathymus yuccae
(Bdv. & LeC.) and its sub species. Ent News 54: 211-217.
1950. Notes on Megathymus, with description of a new species.
Field & Lab. 18: 144-146.
— 1952a. Notes on Megathymus yuccae (Bdv. & LeC.), with descrip-
tion of a new subspecies. Field & Lab. 20: 29-33.
— — ■ — ^ — - 1952b. Two new species of Megathymus. Amer. Mus. Novitates
no. 1953: 1-9, figs. 1-13.
— 1955. Four new species of Megathymus. Amer. Mus. Novitates,
no. 1711: 1-20, figs. 1-34.
1958. A revision of the genera of the Megathymidae, with the
description of three new genera. Lepid. News. 12: 81-92.
I960. Notes on Agathymus in Texas, and the description of a new
species from Mexico (Megathymidae). Journ. Lepid. Soc. 14: 58-62.
1962. A new species of Agathymus from Texas. Amer. Mus. Nov-
itates, no. 2097, 1-7, figs. 1-6.
HARBISON, CHARLES F., 1957. A new species of Megathymus from Baja
California, Mexico. Trans. San Diego Soc. Nat. Hist. 12: 231-262, pis.
18-21.
HOLLAND, W. J., 1930. New species and varieties of North American but-
terflies. Ann. Carnegie Mus. 19: 155-160.
MURRAY- A ARON, E., 1942. Herbert Morrison in Mexico. Fnt. News
53: 142-143.
POLING, OTHO C., 1902. A new Megathymus from Arizona. Ent. News
13: 97-98, 1 pi.
RILEY, CHAS. V., 1876. Notes on the Yucca Borer, Megathymus yuccae
(Bdv. & LeC.). Trans. Acad. Sci. St. Louis 3: 323-343, figs. 25-31.
— • 1877. Additional notes on Megathymus yuccae. Trans. Acad. Sci.
St. Louis 3: 566-568.
SKINNER, HENRY, 1895. A new AEgiale {Megathymus). Can. Ent. 27: 179.
1911. The larger Boreal American Hesperiidae including Eudamus,
Erycides, Pyrrhopyge and Megathymus. Trans. Amer. Ent. Soc. 37: 169-
209-, pi. 10.
STALLINGS, DON B., & J. R. TURNER, 1954. Notes on Megathymus
neumoegeni, with description of a new species. Lepid. News 8: 77-87.
— 1956a. Description of a new subspecies of the Megathymus yuccae
(Bdv. & LeC.) complex. Bull. So. Calif. Acad. Sci. 55: 150-152, pi. 3.
— — ■ 1956b. Notes on Megathymus ursus, with description of a related
new species. Lepid. News 10: 1-8.
— — — 1957. Four new species of Megathymus. Ent. News 68: 1-17.
— — - 1958. A review of the Megathymidae of Mexico, with a synopsis
of the classification of the family. Lepid News 11: 113-137, 8 pis.
— — — — i960. A new species of Agathymus and a new subspecies of Meg-
athymus. Ent. News 71: 109-115.
1961. A new subspecies of Agathymus mariae from Mexico. Journ.
Lepid Soc. 15: 19-22.
STRECKER, HERMAN, 1871. Description of a new species of AEgiale and
notes on some other species of North American Lepidoptera. Proc. Acad.
Nat. Sci. Philadelphia 1876: 148-153.
Skipper from Arizona. Bull. So. Calif. Acad. Sci. 53: 75-87, pi. 2.
TINKHAM, ERNEST R., 1954. The biology and description of a new Giant
Skipper from Arizona. Bull. So. Calif. Acad. Sci. 53: GE-RG, pi. B.
2(2):137-14h
MEGATHYMIDAE
139
TABLE 1
AEGIALE Felder
1, hesperiaris (Walker), vie. Mexico City, D, F. , Mexico
A GA THYMUS Freeman
I. neumoegeni (Edwards), approx. 9 mi. S, Prescott, Ariz.
Z. c a r 1 s b a d e n s i s (Stallings and Turner), Guadeloupe Mtns. ,
Carlsbad Cavern National Park, N. M, , on the Mesa at the
head of Yucca Canyon, elevation 5470 fe et, pH 7, 9.
3. florenceae (Stallings and Turner). Davis Mtns., Scenic
Drive, Texas, elev.-6Z00 ft. , pH 5. 9.
4. judithae ( Stallings and Turner), Hueco Mtns, , approx. 8 mi.
E. Hueco, Texas, el. 5300 ft. , pH 7. 3.
5. diabloensis Freeman, Diablo Mtns. , approx, 5 mi, W. Victoria
Canyon, Texas, el, 5700 ft, , pH 7, 4,
6. mcalpinei (Freeman), 5, 1 mi, N. Marathon, Texas, flats
near foothills of Glass Mtns. , elev. 4300 ft, , pH 7. 4,
7. chisoensis (Freeman), Chisos Basin, Texas, elev. 5400ft, pH 5. Z.
8. juliae (Stallings and Turner), north of Zarca, Durango, Mexico,
on highway 45 at Kim. 1317, elev. 6300 ft,
9. hoffmanni (Freeman), Valle de Mexico, Mexico,
10. evansi (Freeman), Ramsey Canyon, Arizona.
II. belli (Freeman), La Bequilla, Durango, Mexico.
IZ. aryxna (Dyar), The accepted type of this species is figured
in the Biologia Cent. Amer, Lep, Het, III, pi, 69, fig, 4, It
represents very well the maculation of the specimens found
in the Santa Rita Mtns, , Santa Catalina Mtns. , on down to-
wards Nogales, Ariz, Murray-Aaron (Ent, News 53: 143)
states that he and Morrison collected around Nogales and on
down to Hermosillo, Sonora, Mexico, The specimen is label-
led N. Sonora, Mexico, Morrison, indicating that the specimen
was collected in the general vicinity south of Nogales, Sonora,
since Hermosillo is in the central part of the state. We do not
know whether the specimen in question was collected on that
trip or another that Morrison made by himself; the dates of
their collecting trip has been lost. There is some question
as to the status of drucei, which was named from fig, 3 in
the Biologia by Skinner, One specimen from the Chiricahxia
Mtns, and one from Ramsey Canyon, Huachuca Mtns, , Ariz.
are identical with the figure of drucei. Specimens from those
two areas deviate slightly from those from the Santa Rita
and Santa Catalina Mtns, I would like to indicate the type
locality of aryxna as being the western slopes of the Patagonia
Mtns, , S. E» of Nogales, Sonora, Mexico.
13, baueri (Stallings and Turner), Verde Hot Springs, Yavapai
county, Arizona, elev, 4000 ft,
14, freeman! Stallings and Turner, nr, Bagdad, Yavapai county,
Arizona, elev. 5000 ft.
140
FREEMAN
). Res. Lepid.
15, field! Freeman, Guadalajara, Mexico, Jalisco Highway, 1 5,
klm. 724, elev. 4400 ft,
16, mariae (Barnes and Benjamin), Franklin Mtns, , El Paso,
Texas, approx, elev, 3900 ft, , pH 8, 4,
17, micheneri Stallings, Turner and Stallings, approx, 15 mi.
S, Allende, Coahuila, Mexico, elev, 1300 ft, , pH 7, 0.
18, Stephens! (Skinner), Mason Valley (La Puerta), San Diego
coxinty, Calif,
19, c o m s t o c k i (Harbison), 2 mi, N,E, San Simon, Baja Calif, , Mex,
20, remington! (Stallings and Turner), mountains S, of Jacala,
Hidalgo, Mexico, on highway 85, klm, 250, elev, 6000 ft,
21, estelleae (Stallings and Turner), plains 56 mi, S,E, Reynosa,
Mexico, near General Bravo, Nuevo Leon, elev, 400 ft, pH 7, 3.
22, poling i (Skinner), Baboquivari Mtns,, Pima county, Arizona
23, alliae (Stallings and Turner), 15 mi, W, Cameron, Arizona,
along canyon of Little Colorado R, , elev, 5000 ft,
24, indeci sa (Butler and Druce), Costa Rica
25, rethon (Dyar), Sierra de Guerrero, Mexico
TURNERINA Freeman
1, mejicanus (Bell), Guanacevi, Durango, Mexico
2, hazelae (Stallings and Turner), near Chilpancingo, Guerrero,
Mexico, highway 95, klm. 235, elev, 2300 ft,
MEGATHYMUS Scudder
1, yucca e (Boisduval and LeConte)
a, yucca e (Boisduval and LeConte), Aiken county. South
Carolina is here designated type locality as it is the
nearest place to Abbot's home in Scriven county, Georgia
where he is known to have collected, and where Yucca
grows. Yucca smalliana and Y, flaccida,
b, buchholzi Freeman, Jupiter, Florida. Yucca glorios
c, stallings! Freeman, Caldwell, Kansas, pH 6, 1, Yucca
arkansana,
d, wilsonorum Stallings and Turner, Victoria, Tamalipas,
Mexico. Yucca treculeana and Y.carnerosana
e, coloradensis Riley, vicinity of Colorado Springs, Colo,
Yucca glauca,
f, navajo Skinner, Ft, Wingate, Zuni Mtns,, New Mexico,
Yucca baccata,
g, arizonae Tinkham, Mountain View, Pima county,
Arizona, Yucca elata and Y , Thornberi,
h, martini Stallings and Turner, Little Rock, Los Ang-
eles county, Calif, Yucca brevifolia,
i, browni Stallings and Turner, Salina, Utah, Yucca
harrimaniae,
2. cofaqui (Strecker), The allotype was collected at Boca
Grande, Florida which is here designated as the type locality.
The exact locality of origin of the holotype is believed to
2(2);1}7-141, 1963
MEGATHYMIDAE
141
be somewhere in northern Florida near Georgia, collected
by Morrison, Yucca aloifolia,
3, harrisi Freeman, Stone Mountain, Georgia, Y u c c a
filamentosa.
4, streckeri (Skinner), Type locality here designated at
Petrified Forest area, Arizona as photos of the type
agrees best with material from that area. Yucca
Standleyi,. probably,
5, texanus, Barnes and McDunnough
a, texanus Barnes and McDxinnough, 2d'cf,29$ collected
by Jacob Boll at Dallas and San Antonio, Texas, To
this collector Dallas was North Texas and San Antonio
was South Texas, Kerrville, Texas is here desig-
nated at the type locality by reason of being the
nearest known’place to the above where texanus
may be found. Yucca glauca, associate,
b, leussleri Holland, Sand Hills near Valentine, Neb-
raska, Yucca glauca, probable,
6, ursus Poling, Santa Catalina Mtns, , west of Redington,
Pima county, Arizona, pH6.1. Yucca Schottii.
7, viola e Stallings and Turner, Carlsbad Cavern National
Park, New Mexico, elev, 4700 ft, , pH 7, 5, Yucca Torreyi.
8, beulahae Stallings and Turner, near Ixmiquilpan, Hidalgo,
Mexico, highway 85, klm, 176, elev, 5700 ft. Agave ? near
Schottii,
STALLINGSLA Freeman
1, smithi (Druce), Amula, Guerrero, Mexico, approx, 3000 ft,
Manfreda maculata, probable,
2, rhac.ulosus (Freeman), 2 mi, S, Kingville, Texas, elev,
100 ft,, pH 7. 0, Manfred maculosa.
142
FREEMAN
/. Res. Lepid.
BIOGRAPHICAL SKETCHES
FREEMAN, HUGH AVERY
[1605 Lewis Drive, Garland, Texas]
Born; Conway, Arkansas, Oct* 7, 1912
Married; 1939, 3 children
A„ B, ; Hendrix College, 1936
M. S, ; Southern Methodist University, 1938
Teacher; High school Texas, 1938-48
Instructor; Biology, Southern Methodist University, 1948-51
Teacher; High school, Dallas, 1951
Interests; Started collecting lepidoptera in 1928
Began specializing in the hesperi -
oidea in 1938, In 194Z started in-
tensive study of the Megathymidae,
which is still going on. Also inter-
ested in the study of metal marks
and hair streaks. Has published 44
articles on lepidoptera, most of
which were upon the hesperioidea.
Journal of Research on the Lepidoptera
2(2) :143-150, 1963
1140 W. Orange Grove Ave., Arcaiia, CaUforms, US. A.
© Copyrigbi 1^63
THE DISTRIBUTION OF
AN ENDEMIC BUTTERFLY
LYCAENA HERMES
FRED THORNE
1360 Merritt Drive
El Cajon, California
One of the most interesting of the endemic butterflies is
Lycaena hermes (Edw.). Its known range extends fifty miles north
of the Mexican border almost to Failbrook, San Diego County, Cali-
fornia, and south of the border aimos; one hundred miles to a point
eighteen miles south of Santo Tomas, Baja, California, Mexico. In
San Diego County it ranges inland to Pine alley, about forty miles
from the Pacific Ocean.
Older literature and check-lists tended to overstate the range, i. e.,
California and Nevada, whereas all recently published information
understates the range. In the light of present knowledge, the area
occupied may be as large as the state of Connecticut. However, within
this range its distribution is limited to pockets where the larval
food plant occurs, so that the total area where the insect acually flies
is probably not more than a fraction of one percent of the maximum
area. Such limited distribution within a given range is not unique
with endemics, being a common occurence among plants and animals.
Colonies of the Hermes Copper are closely confined to the vicinity
of the host plant, Rhamnus crocea Nutt. Extensive collecting for over
thirty years has failed to produce specimens beyond a short distance
from the larval food plant. There is no observable tendency to migrate,
to "hilltop,” or otherwise to stray from these colonies, although there
must be some inter-colony movement, probably by the males. Popula-
tions within the known range therefore depend on the distribution of
the host plant, and there is. certainly nothing novel about this among
insects.
It is very difficult to analyse the complex factors- which determine
why a certain plant has been successful in a 'given spot, and why
it has been able to out-compete all other plants for this particular
place in the sun. In the case of Rhamnus crocea, the only consistent
requirement seems to, be a well drained soil of better than average
depth, yet not deep enough to support trees. Such soils occur along
canyon bottoms and on hillsides with a northern exposure; therefore,
it is in these, situations that hemes is -generally found.
143
144
THORNE
). Res. LepiJ.
A notable point about hermes is that the host plant extends well
beyond the range of this insect, Rhamnus crocea, in one form or another,
extends to Mt. Diablo in the coastal ranges of California; along the
foothills of the Sierra Nevada; on the Channel (Santa Barbara) Islands;
and even into the Mojave Desert. Yet within its range in San Diego
County, hermes tolerates greater climatic extremes • than it would en-
counter in some of the contiguous coastal areas open to colonization,
and in insular areas which might have been available at one time.
Freezing winters with snow are normal in Pine Valley at an
elevation of 3800 feet, while summer temperatures of 105 °F. are not
uncommon in some of the foothill localities. To borrow an appropriate
sentence from Flovanitz (1963), said of the distribution of Argynnis
idalia, "The biological reasons for this restricted distributional range
are not known." Biologists will appreciate how often this same con-
clusion must be drawn.
Nelson (1921) recognized a San Diegan Faunal District in north-
western Baja California which roughly corresponds with the area
occupied by hermes in that Mexican state, but did not define the
northern limits of this life zone in Southern California. A few species
of plants appear to be restricted to about the area inhabited by
hermes, but there is little to suggest, at least north of the border, that
the area should be segregated from the Upper and Lower Sonoran
life zones usually assigned to it. Nor has northwestern Baja California
been shown to be an originator of new species, since the coastal fauna
shows a close affinity to that of coastal Southern California, and the
mountain fauna is simply an extension of the southern Sierran with
the exception of a few instrusions (Rindge, 1948; Powell, 1958; Pat-
terson and Powell, 1959; Truxal, I960).
L. hermes is in a good state of balance in its environment. The
season of emergence for the adults is very dependable, as is their
presence every year in their select habitats. There is no wide fluctuation
in numbers from year to year, although the current prolonged drought
has reduced the populations in common with nearly all Lepidoptera.
It would be difficult to believe that it is not autochthonous. Fossil
evidence of insect distribution is so limited that it will probably
never be known whether hermes ranged over a wider territory than
now. It is an insect which seems to exhibit stability due to long occupa-
tion of its present habitat, yet it is difficult in the light of other biologi-
cal evidence to view the present range as a refugium of some sort.
Concerning the genus Lycaena, Clench (1961) has stated, "A
curious and possibly quite ancient genus, strongly developed in both
the Palaearctic and the Nearctic regions, with a small handful of
outliers — one in Guatemala, one in South Africa and several, most
perplexing, in New Zealand.’’ Perhaps this offers a clue, yet careful
studies by Klots (1936) and Freeman (1936) have indicated that
2(2):143-1^0, 1963
DISTRIBUTION LYCAENA HERMES
145
hermes has affinities with other North American members of the
genus. Kiots says, "Evidently hermes is, structurally at least, far closer
to gorgon and heteronea than to its tailed Nearctic congeners." Free-
man placed hermes in his xanthoides group on the basis of the genitalia,
together with dione, rubidus, and editha.
Another interesting thing about hermes is the use of a species of
Rhamnaceae as the larval host. Most congeners feed on Polygonaceae
(Eriogonum, Rumex, Polygomim)\ some of Rosaceae {Potentilla) ]
some on Saxifragaceae {Ribes)\ and some on Ericaceae {Vaccinium),
but this use of a Rhamnaceous plant is believed to be unique for
the genus (Davenport and Dethier, 1937). The commonest source
of nectar for the imagines is Eriogonum fasciculatum Benth., one of
the Polygonaceae, and this plant is almost invariably present. Could
this in some ancient time have served as the larval host?
One might reason that the failure of hermes to invade large areas
which appear to be open to colonization might be due to prior occupa-
tion of particular ecological niches by another species (competitive
exclusion). The unusual food plant which is not used in other areas
by any member of the genus would seem to rule this out. Or, if one
wishes to consider hermes as the victim of some complex, predator-
parasite relationship which grew up around one of the congeners
and favored it over hermes, Lycaena {T harsalea) arota would be the
most logical candidate. This species appears to be at the extreme
southern limit of its range in San Diego County as evidenced by the
few specimens ever taken. Since it feeds on Ribes, there is no direct
competition, and the hypothesis becomes even more dubious from
the fact that hermes lives sympatrically, or at least there are zones of
contact along watercourses with Lycaena helloides and L. xanthoides.
These should reasonably be expected to furnish whatever environ-
mental pressures that arota might.
There is rather general belief that hermes is in a last ditch struggle
for survival in San Diego County. This isn’t true! Colonies have
survived in areas that have been overrun with houses for many
years; in areas being grazed by livestock; in areas being farmed
(avocado orchards); and in areas which have been burned-over with
some frequency. The map, Fig. 1, shows the wide distribution of
known colonies which should ensure survival for the foreseeable
future. This map should not be regarded as a complete record of
distribution, since accessibility by road has been the main factor in
locating colonies.
The insect has been beautifully illustrated in Comstock’s "Butter-
flies of California” and his poetic description of the butterfly in
nature is worth repeating. He says, "It is a fascinating little sprite as
it darts about in the sunlight, or sports is showy colors while balanced
on a tuft of wild buckwheat." As for the flight period, my earliest
146
THORNE
J. Res. LepiJ.
record is May 20, 1934, and the latest is July 20 at Alpine by George
Field. Records from several hundred captures show peak flight about
June 20, but the best time to collect males is about June 10, and
for the females about June 20. Field captures show a large preponder-
ance of males (85%), but this a probably a false indication of the
actual sex ratio because of the mo’-e retiring habits of the females, and
because of their tendency to flee directly from the place of disturbance
so that they are quickly lost to sight. However, the percentage of
females increases late in the flight period in common with many
butterflies. The males practice territorialism, but are not very aggre-
sive about it. They will patrol a section of flyway, or watch it from
a vantage point, often on the host plant, but from any suitable perch.
Both sexes visit flowers avidly, and the blossoms of Eriogonum fasci-
culatum supply the bulk of nectar.
The species is single-brooded and spends about two-thirds of its
life in the egg stage. It aestivates and hibernates in the ovum, and the
hatching of one egg was observed under field conditions on March 16.
Mature larvae were recovered by beating the host plant on May 24
near Lyons Peak, where the season is delayed due to elevation. The
egg, mature larva, and pupa have been illustrated by Comstock and
Dammers ( 1935). Females oviposit readily in captivity, but unless
the ova are kept on a living plant, they fail to hatch. The reason for
this is not known, but may simply be dessication. Nothing in this
life history sets hermes apart in any remarkable way from other mem-
bers of the genus, although several are multivoltine.
As a generalization, most endemics are univoltine (consider the
alpine relicts) but there are numerous exceptions. A good example is
Strymon avalona Wright, another interesting endemic, which is con-
fined to Santa Catalina Island off the southern California coast despite
the fact that its food plant, Hosackia (Lotus) argophylla Gray, is more
widespread. This insect has a succession of broods. The biological
reasons for its restricted distributional range are not known, but are
easier to deal with than is the case with hermes, since insular, alpine,
acid bog, or other types of endemism offer the biologist some
solace during the brief instant in evolutionary time that he is around
to observe distribution.
Populations of the Hermes Copper in each colony are not great,
probably numbering in the hundreds. Six sample counts taken at
random from field notes for 1955 to 1959 show the capture of 69
specimens in 405 minutes, about 6 minutes per catch. Any day in
which 50 specimens are taken can be regarded as exceptional. It is
entirely fair to regard the insect as "not uncommon" as expressed by
Clench (1961) — in fact, it falls comfortably into Clark’s (1932)
standard of "abundant” (where fifteen or more can be taken in an
average day ) .
2(2):14}-n0, 196}
DISTRIBUTION LYCAENA HERMES
147
, PAL OMA R M TN.
ST ATE PARK
O
Fig. 1. Map showing the distribution of Lycaena hermes in San Diego
county, California. . i
148
THORNE
). Res. Lepid.
It has been stated previously that there must be some inter-colony
movement. The basis for this is that differences among the populations
are not readily observable, if indeed any exist. It must be admitted
that no effort has been made to comoare adequate series from
different colonies to see if any segregation is evident. This vv^ould be an
interesting study, but until it is made, it seems best to assume that
gene flow throughout the entire range is adequate to prevent segregation.
If this is true, then the gene pool has some magnitude. Nevertheless,
it seems likely that hermes is a conservative species, homozygous for
many of its characters, variability being restricted by a close adaptation
to a narrow environment. Barriers to spread appear to be intrinsic,
that is, because of the inherited behavior patterns, the imagines "choose
to remain” within very limited areas despite their ability to fly else-
where, and presumably, to occupy larger territories (see Ehrlich, 1961).
The chromosome number has not been published yet (N=24 for most
species of the genus (Maeki and Remington, I960) but this infor-
mation is expected to be available soon.
Perhaps the distinctive facies and unusual food plant of hermes
represent a genetic breakthrough which will result in time in a more
widespread and successful species. Only time will tell whether the
insect has retreated into a final refugium, the nature of which is not
too evident, or whether it has the genetic resources eventually to
expand its habitats.
That such expansion is possible has been demonstrated in a
spectacular way by Paratrytone melane melane (Edw.) which was
not recorded from San Diego County prior to 1941. Wright (1930)
said, "Further collecting in wooded areas of the county may produce
this species.” Gunder (1930), in his now classic checklist of the
butterflies of Los Angeles County wrote, "Never seemingly abundant
in one locality, but may be had, several at a time each year.” Rindge
(1948) recorded this insect from southern Lower California, but
this is now regarded as a distinct and undescribed sub-species (Mac
Neill, 1962).
I first encountered this skipper on July 20, 1941, when two speci-
mens were collected near El Cajon, California. Visits to the same
spot July 23, 24, 25 and August 2 and 3 resulted in fifteen more
specimens, and show my interest in what I thought was a once-in-a-
lifetime chance. Other collectors also reported finding this species for
the first time in San Diego that year. On October 5, 1941, a specimen
was taken in the desert at Mason Valley, San Diego County. Since
then, quite contrary to my expectations, this insect has become one
of the very common skippers in my garden, flying from February
to December. Powell (1958) records it from northwestern Baja,
California in what I believe to be a further extension of this same
population explosion.
2(2):l4}-n0, 196)
DISTRIBUTION LYCAENA HERMES
149
This is an example of an insect which suddenly expanded into
and occupied contiguous areas which are evidently well suited to
it. It is not the purpose of this article to discuss this interesting
phenomenon, but simply to point out that extensions of range are
possible, and that it may be the privilege of the lepidopterist to see
it happen in his own yard.
SUMMARY
The known range of Lycaena hermes (Edw. ) extends from fifty
miles north of the Mexican border in San Diego County to one
hundred miles south of the border in Baja California. The insect occurs
in colonies around the foodplant Rhamnus crocea, but has failed to
invade other areas that appear suitable for reasons that are not known.
The life history and field behavior are not unusual. The species is
believed to be autochthonous and conservative, perhaps in a final
refugium, but spread to other areas is possible, as has been demon-
strated by Paratrytone melane (Edw.)
REFERENCES
BROWN, F. MARTIN 1962. Notes about the types of some species of
butterflies described by William Henry Edwards. Ent. News. 73 (10):
265-268 (pg. 267).
CLARK, AUSTIN H. 1932. Butterflies of the District of Columbia. Smith-
sonian Inst. U. S. Nat. Mus. Bull. 137, Wash. D. C. (pg. 25),
CLENCH, HARRY K. 1961. In, How to know the butterflies. Ehrlich and
Ehrlich. Brown Co., Dubuque, Iowa (pg. 22).
COMSTOCK, JOHN A. 1927. Butterflies of California. Los Angeles, Calif.
(pg. 172 and PI 51, Figs. 9-11).
COMSTOCK, JOHN A. and CHARLES M. DAMMERS 1935. Notes on the
early stages of three butterflies and six moths from California. Bull. So.
Calif. Acad. Sci. 34 (2); 120-141. (pp. 124-126).
DAVENPORT, D and V. G. DETHIER 1937. Bibliography of the described
life histories of Rhopalocera of America north of Mexico 1889-1937.
Ent. Amer. 17 (New series) (4); 155-196. (pp. 174-175).
DRAUDT, Dr. M. 1924. In Seitz, Macrolepidoptera of the world Vol. 5.
American Rhopalocera. Stuttgart, Germany, (p. 812).
EDWARDS, WILLIAM HENRY 1870. Description of new species of Lepi-
doptera found within the United States. Trans. Amer. Ent. Soc. 3:21
original description of Chrysophanus hermes.)
EHRLICH, PAUL R. 1961. Instrinsic barriers to dispersal in checkerspot
butterfly. Science 134, No. 3472: 108-109.
FREEMAN, T. N. 1936. Notes on the specific grouping of the genus Lycaena
( Lepidoptera ) . Can. Ent. 68: 277-279.
GUNDER, J. D. 1930. Butterflies of Los Angeles County, California. Bull.
So. Calif. Acad. Sci. 29 (2): 1-59 (on page 58).
HOLLAND, W. J. 1931. Butterfly Book (Revised ed.), Doubleday and Co..
New York, (on page 247).
HOVANITZ, WILLIAM 1963. Geographical distribution and variation of
the genus Argynnis. J. Res. Lepid. 1 (2); 117-123 (on page 123).
JEPSON, WILLIS LINN 1925. Manual of the flowering plants of California.
Univ. of Calif. Press, Berkeley, Calif, (on pages 614-615).
150
THORNE
}. Res. Lepid.
KLOTS, ALEXANDER B. 1936. The interrelationship of the species of the
genus Lycaena Eabricius (Lepid., Lycaenidae), Bull. Brooklyn Ent. Soc.
31 (4); 154-171. (on page 164).
MacNEILL, C. DON 1962. A preliminary report of the Hesperiidae of Baja
California. Proc. Calif. Acad. Sci., 4th series, 30 (5): 91-116.
MAEKI, KODO and CHARLES L. REMINGTON I960. Studies of the
chromosomes of North American Rhopalocera, Part 3. /• Lep. Soc. 14 (2) :
127-147.
MUNZ, PHILIP A. 1935. Manual of Southern California Botany, Claremont
Colleges, Claremont, Calif, (on page 299).
NELSON, EDWARD W. 1921. Lower California and its natural resources.
Mem. Nat. Acad. Sci., Wash., D. C. 16: 1-194. (on page 118).
PATTERSON, DONALD and JERRY A. POWELL 1959. Lepidoptera col-
lecting in the Sierra San Pedro Martir, Baja California J. Lep. Soc. 13:
229-235.
POWELL, JERRY A. 1958. Additions to the knowledge of the butterfly fauna
of Baja California Norte. Lep. News 12: 26-32.
RINDGE, FREDERICK H. 1948. Contributions toward a knowledge of the
insect fauna of Lower California. No. 8 Lepidoptera; Rhopalocera. Proc.
Calif. Acad. Sci., 24: 289-311.
TILDEN, J. W. 1955. A revision of Tharsalea Scud. (s. str.) with descrip-
tion of a new subspecies (Lepid., Lyc. ) Bull. So. Calif. Acad. Sci. 54
(2): 67-77.
TRUXALL, FRED S. I960. Symposium: Biogeography of Baja California
and adjacent seas. The entomofauna with special reference to its origins
and affinities. Syst. Zoo. 9 (1-4): 165-170.
WRIGHT, WILLIAM GREENWOOD 1906. Butterflies of the West Coast.
Second ed. San Bernardino, Calif.
WRIGHT, WILLIAM S. 1930. Annotated list of the butterflies of San Diego
County, California. Trans. San Diego Soc. Nat. Hist. 6 (1): 1-40 (on
pages 25, 26 and 40).
Journal of- Research on the Lepidoptera
2(2) :151-160, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright
CALLOPHRYS (LYCAENIDAE) FROM
THE PACIFIC NORTHWEST
HARRY K. CLENCH
Carnegie Museum, Pittsburgh, 13, Pa.
Over two years ago Mr. E. J. Newcomer, of Yakima, Wash-
ington, sent for examination and identification a series of Callophrys
{Callophrys) taken near his home. He subsequently augmented this
lot by other specimens, some of- which were also taken locally while
others were from farther afield in the state. More recently Mr. David
L. Bauer, of Bijou, California, has sent for study additional specimens
of this subgenus from both Washington and Oregon. The present
paper is concerned chiefly with this assembled material. My best
thanks are due both these gentlemen for the opportunity to study
and report on this now sizable and instructive collection from a region
whence before Callophrys had been all but unknown.
Callophrys is in several ways a very difficult subgenus. Problems
of discriminating species occur in several regions, notably (but by no
means exclusively) that under discussion in this paper. An even greater
problem exists in the group as a whole: the correct association in
polytypic species of the various named entities. Six species are cur-
rently recognized (Clench, 1961) in the subgenus {dtimetorurn- Bois-
duval, corns tocki Henne, apama Edwards, af finis Edwards, viridis
Edwards and sheridanii Edwards ) . a number which is probably too
large. On the other hand^ we know that there must be at least two
species for two occur sympatrically in many places: sheridanii and
apama in Colorado; sheridanii and af finis in the Yellowstone area
and near Brewster, Washington, sheridanii and dumetornm near
Yakima, Washington; viridis and dumetornm in the San Francisco Bay
area. But throughout their known ranges apama and affinis do not;
nor do affinis and dumetornm; apama and dumetorum; sheridanii and
viridis; or affinis and viridis.
It is tempting, and it may possibly be correct, to associate sheridanii
and viridis as one species; dumetorum, comstocki (which is sympatric
with no other), apama and affinis as another. This arrangement would
satisfy nicely the conditions of sympatry, but it creates other problems;
and besides, it is sheer speculation.
The traditional method of resolving such difficulties is by study
of the male genitalia. Barnes and Benjamin (1923), the first authors
to revise the group, did not make use of these structures nor did I
151
152
CLENCH
). Res. Lepid.
in my revision of some years ago (Clench, 1944). The present occasion
seemed an excellent opportunity to fill this long-standing desideratum.
Preparations were accordingly made of all of the major and most of
the minor entities, usually of several individuals each, and these prep-
arations subjected to study. Preliminary examination failed to disclose
any qualitative differences but did reveal the existence of considerable
variation in relative lengths of different structures. A series of measure-
ments was then made and these carefully and elaborately analyzed
individually, paired in correlations, as ratios and by other more esoteric
means, which it is pointless to describe as the results of the whole
inquiry were thoroughly, discouragingly negative: the male genitalia of
Callophrys s.s. show considerable individual variability but give no
evidence whatever of variation useful for discriminating species. We
are still where we were before.
Three species of Callophrys s.s. are now known from the Pacific
Northwest^ one here recorded for the first time. They epitomize the
difficulties noted just above and it would be fatuous to presume that
the results here given represent the final word on the solution of those
difficulties. Decisions on relationships had to be made and I have
tried to make them as accurately as possible.
Callophrys {Callophrys) af finis washingtonia Clench
Known from Alta Lake (type locality) and Brewster, Washington
and from Summerland, Osoyoos, Penticton and Keremeos, British
Columbia, the latter records in the Canadian National collection.
Size rather large (about as in dumetorum perplexa or apama)\
ground color above greyish with a distinct brownish tinge and in the
female with some discal fulvous (occasionally in the male as well);
underside apple green, a little less yellowish than in nominate affinis
but decidely yellower than other members of the genus; on the fore-
wing the green extends posteriorly to about CU2, thus covering most
of the wing; postmedian spot row completely absent or at most repre-
sented by faint, hardly visible traces. Base of fore wing costa below
pale grey.
Callophrys {Callophrys) dumetorum Boisduval
When Mr. Newcomer sent his first series of Callophrys he wrote
that he was interested to learn whether they represented one species
or two, for some were immaculate below like affinis, others well
marked, more or less as in sheridanii. Study showed that all of this
variability actually was comprised in the one new subspecies of
sheridanii described below. There was a surprise for us both, how-
ever, that came to light only when the specimens were being studied
after spreading: a single fresh male of dumetorum from Satus Creek,
2000 ft., Yakima Co.^ Washington, taken in 16 May I960. When I
wrote to him about this discovery and described its' differentiating
characters, Mr. Newcomer checked through his material and found
an additional specimen from the same locality taken I6 May 1959.
Returning to the same area in the spring of 1961 he was able to
2(2):151-I60, 1963
CALLOPHRYS FROM NORTHWEST
153
take several additional specimens most of which he sent on for
study: Kussdhi Creek, off Sams Creek, 2000 ft., Yakima Co., 24
May 1961 (1 female), and 2 June 1961 (1 male); Ski area near
Sams Pass, 3700 ft., Klickitat Co., 2 June 1961 (1 female). In the
material sent by Mr. Bauer were two males of this species from
Shelton, Mason Co., Washington, 2 May 1958. All these specimens
represent a considerable northward extension of the known range of
dumetofum (cf. Clench, 1961; 210).
Three regional lists relating to the area under consideration in this
paper have had to be ignored: Bowman (1919) on Alberta; Blackmore
(1927) on British Columbia; and Leighton (1946) on Washington state.
Each lists dumetomm and no other Callophrys s. s. (except Leighton, who
also lists dumetomm perplexa and sheridanii) , misdeterminations (partial
or complete) rather typical of the trouble these butterflies have given to
systematists.
Males [Description refers only to Washington material.] are uniform
grey above with a very slight brownish cast. Females above are
largely dark fulvous, shading to brownish fuscous in the basal third
of both wings, along costa, termen and (more broadly) apex of fore
wing as well as very narrowly along termen of hind wing. The fringe
is greyish white (markedly duller than the fringe of either af finis or
dumetomm), darker basad. On the under surface of both sexes the
inner marginal grey of fore wing extends costad usually to M3; the
postmedian line of this wing is tolerably well developed from Cuq to
Ml or R4, but often rather faint, occasionally even obsolete (though
a trace of the dark basal part of the line usually persists ) . The post-
median line of the hind wing is highly variable in its expression, rang-
ing from nearly complete (bars in ail interspaces from Sc to inner
margin) to nearly absent (slight whitish bars in Sc - Rs and Cui -
CU2); base of fore wing costa below usually fulvous. Size rather small:
about the same as s. sheridanii.
The subspecies of dumetomm have never been worked out adequa-
tely. Even the range of the species is imperfectly known. For the
present these Washington specimens seem best left under nominate
dumetomm with which they agree far more closely than with d.
perplexa Barnes and Benjamin from the lowlands of southern Cali-
fornia.
Callophrys {Callophrys) sheridanii neoperplexa Barnes and Benjamin
The upper surfaces of both sexes are uniform grey with no
tinge of brownish or fulvous. Base of fore wing costa below pale
grey. Under surface green, on fore wing extending posteriorly to
Cu2; postmedian lines well developed on both wings, usually more
or less continuous (the component white bars conjoined), but rather
thin and lacking the black basal edging characteristic of s. sheridanii.
Size small.
In the Pacific Northwest known only from Brewester, Washington
where the late J. C. Hopfinger used to take it rather frequently. Until
154
CLENCH
). Res. Lepid.
the receipt of Mr. Newcomer’s material it was not realized just how
unusual this record is. See the discussion below.
Callophrys ( Callophrys ) sheridanii newcomeri new subspecies
The upper surfaces of both sexes are uniform grey, unrelieved by
any brownish tint or fulvous, much as in the other sheridanii sub-
species. Base of fore wing costa below, pale grey. It differs chiefly in
the strong tendency to reduction of the postmiedian line of the under-
side, continuing the trend away from s. sheridanii that is evident in
s. neoperplexa Barnes and Benjamin. This is especially marked on
the fore wing where most specimens show either no trace of the
line at all or only a few faint bars; occasionally it may be fully
present but is then only faintly developed. In only one specimen, a
female from Ft. Simcoe, is it fully present and strongly developed.
The row on the hind wing is rarely completely absent (a female
from Mill Creek), but almost always lacks several bars at least,
especially those in Rs - Mi and M3 - Cui. On the fore wing upper
side of the Mill Creek series there is a definite, though faint, pale
grey patch, strong at cell-end and fading rapidly distad.
Holotype, male, and 5 male paratypes, all Mill Creek, 1800 ft.,
Yakima Co., Washington, 29 March 1961 (E. J. Newcomer) the
holotype including male genitalia slide C-791; 5 male and 7 female
paratypes, Ft. Simcoe, 1200 ft., Yakima Co., Washington, 6 April I960
(E. J. Newcomer).
Holotype and most of the paratypes, C. M. Ent. type series no.
478. Some paratypes are being returned to Mr Newcomer.
Remarks In addition to the type series, the following material
has been seen: Mt. Spokane, 5000 ft., Spokane Co., Washington, 27
June I960 (E. J. Newcomer), 3 males and 2 females; Blue Mts.,
4800 ft., Columbia Co., Washington, 17 June 1961 (E. J. Newcomer),
2 males; Lonerock, Gilliam Co., Oregon, 7 June 1961 (D. L. Bauer),
2 males and 2 females (doubtfully typical).
The Canadian National Collection has specimens from Waterton
Lakes Park, Alberta, and from Okanagan Landing and Vernon,
British Columbia, all of which I believe are referable to newcomeri,
though I saw them but briefly while on a visit in 1956 and made only
a few notes on their peculiarities.
The Oregon population and the four from Washington all show
a certain amount of independent differentiation. The greatest departure
is found in the Lonerock series where females show a definite fulvous
tinge above, unique in the species, and the under surface (which is
more uniform in appearance than in any of the Washington popula-
tions) has the postmedian line on both wings practically obsolete save
for the portion posterior to Cui on the hind wing. Should these dif-
ferences hold true in a larger series it may be advisable to separate
this under another subspecific name.
2(2):IU-160, 196}
CALLOPHRYS FROM NORTHWEST
155
The fresh series of males from Mill Creek shows a faint, small,
pale discal patch on the fore wing above as noted above; this also
occurs in some of the males from Ft. Simcoe, though more weakly.
The Mill Creek series also averages somewhat larger than the others.
The specimens from Mt. Spokane are somewhat smaller, tend to
be somewhat darker below and have the postmedian line of the
fore wing below absent completely.
Several points concerning sheridanii in the Northwest merit
further discussion. These points may be grouped under two broad
headings: geographic variation and climatic adaptation.
Geographic variation. The differences between sheridanii popula-
tions from place to place are of a rather unusual nature. The most
conspicuous aspect of this is the isolated occurrence of s. neoperplexa
at Brewster, Washington, remote from the nearest other populations of
that subspecies in southwestern Montana and with much of the
intervening area occupied, apparently by s. newcomeri ( see fig. 1 ) .
It is not yet known whether this Brewster neoperplexa represents an
enclave entirely cut off from other neoperplexa populations or a
peninsula-like intrusion, perhaps along river valleys.
The population of neivcomeri at Lonerock, Oregon, differs more
from any of the Washington populations than these do from each
other, rather surprizing in view of its close geographic proximity.
Lonerock and Ft. Simcoe, for example, are much closer than Ft.
Simcoe is to Mt. Spokane; and both Lonerock and the Blue Mts. locality
in Columbia Co., Washington, are on the same (northwest) slope
of the same mountain range. In addition to its greater differentiation,
the Lonerock series also seems less individually variable. Again it
should be pointed out that the small size of the series makes these
observations tentative and uncertain.
In Washington the known populations of newcomeri show a
definite but rather low level of interpopulation differentiation as
already discussed; and within each population a rather high level of
individual variability. The interpopulational differences seem to bear
little relation to the geographic distances separating the populations,
the Mill Creek and Ft. Simcoe populations, for example, being only
a little less different from one another than either is from the Mt.
Spokane series.
There is evident here a definite hierarchy of differentiation: the
greatest, that between newcomeri and neoperplexa; next, in neivcomeri,
between the Washington populations collectively and the one popula-
tion from Oregon; and finally, the slight differentiation between the
Washington populations. In a general way this hierarchy undoubtedly
reflects the past history of the species in this region and it is a
temptation to draw on it as well as on some of the other data on
variation given above in speculation on the past events that may
156
CLENCH
I. Res. Lct>id.
liave led to the situation as it is today. Until larger series can be
obtained, however, the data basic to such speculation would be far
too shaky. My repeated emphasis of this point is in no way intended
as a slight to the much appreciated efforts of Mr. Newcomer and
Mr. Bauer; their collections are by no means small as series go in
Callophrys and, further, they were not collecting with statistical needs
in mind. Statistics is a notoriously avaricious taskmaster.
Climatic adaptation. The various factors which together make
up what we call climate exert a strong control over the distribution
of Lepidoptera as they do for many other groups of organisms. Indeed,
we may imagine the range of a species of butterfly or moth as an
area bounded by not one but a number of lines, each one representing
a limiting value, for that species, of some particular climatic factor.
Such a notion, evidently, is a great oversimplification but the principle
is valid and useful.
We may touch on the problem only briefly and incompletely
here in connection with C. sheridanii. Two types of climatic responses
may be discerned: (1) responses to particular factors singly, and
( 2 ) responses to two or more jointly ( correlated responses ) . The
data used are weather bureau mean figures as recorded, for example,
in Climate and Man (U.S.D.A. Yearbook of Agriculture, 1941). On
no account is it to be imagined that the insects are necessarily respond-
ing directly to these variables as such. It is merely that they appear
to function as indices more or less closely correlated with whatever
factors may actually be responsible, factors which themselves may be
quite inaccessible for analysis over a large area.
Three variables were selected for study: mean January temperature,
m.ean July temperature, and mean annual precipitation. Values of these
were tabulated for all the known sheridanii localities for which they
were available, as given in Table 1. By inspection we may establish
the approximate limits for each of these;
mean January: all records fall between 14° and 32 °F
mean July: all records fall between 57° and 76° F
ann. precip.: all records fall between 10 and 25 inches
The map ( fig. 1 ) shows the results of applying these limits over the
northern part of the range of sheridanii: all the shaded areas lie out-
side them and hence are presumably unavailable to sheridanii.
Obviously these three factors are not sufficient to account for the
whole of the present range of the species. A large part of Montana
(east of the mountains) and of South Dakota, for example, fall with-
in these limits, yet sheridanii does not occur there. It is quite probable
that variables other than these three are, at least in part, responsible.
Yet without extending observation beyond them the distributional
limits of sheridanii could be approximated still more closely by the
use of correlated responses.
2(2):I)I-I60, /y6.i
CALLOPHRYS FROM NORTHWEST
157
TABLE 1
Mean temperature (® P) for July and January tod mean annual preclpata-
tlon (inches) for localities where Callophrys (Callophrys) sheridanil
has been taken.
Station
Locality
mean
Jan.
mean
Jul.
meui ani
precip,
1
Wash.:
Ft. Simcoe
28.4
75.2
12.29
2
Blue Mts, (Columbia Co.)
28(ca.)
68(ca.)
?
3
Mt. Spokane, 5000 ft.
17.2
58.7
22(ca.)^
4
Brewster
27.6
73.8
10.66
5
Oregon;
Lonerock
31.7
62.0
15.442
6
Mont.:
Dillon
24.4
65.1
16.67
7
Ennis
21.7
64.5
10.69
8
Polaris
20(ca.)
62(ca.)
?3
9
Wyo.:
Centennial
21.5
61.4
16.91
10
Colo.:
Boulder Co,
28(ca.)
71(ca.)
11
Red Feather L., 8400 ft.
15.0
57.9
^4
12
Ft. Collins
26.0
68.9
15.20
13
Utah:
Stockton
25.5(ca.)
72.3(ca.)
12.90^
14
N. Mex.
: Cloudcroft
30.1
59.7
24.58^
1, Temperature values are those of Spokane reduced equally by a lapse
rate of 1° P/300 ft. Precipatation estimated from regional values.
2, Spots for Loneroek^ Oregon, and Cloudcroft, New Mexico, are omitted
from the graph. They fall so far outside the pattern of the
remaining localities that I strongly suspect them of not represent-
ing the particular localities where the sheridanil were actually
tidcen. There is no error in the values themselves which were
kindly confirmed for me by Mr. T. L. Long, National Weather Records
Center, Asheville, North Carolina, Needless to say, this is a
common problem in mountainous country where small differences
in elevation or exposure can exert major changes in the climatic
picture.
3, Estimated values.
4, Temperature values are those of Ft. Collins reduced as described in
note 1,
5, All values are interpolated from those of two adjacent bracketing
localities .
158
CLENCH
/. Res. Lepid.
Fig. 1. Map of a portion of the range of C. {Callophrys) sheridanii
showing localities of subspecies neoperplexa (open circles) and subspecies
neivcomeri (solid circles). Excluded areas (shaded) have one or more of the
following: (1) mean January temperatures below 14°F or above 32°; (2)
mean July temperatures below 57° or above 76°; (3) mean annual precipita-
tion below 10 or above 25 inches.
The method of correlated responses makes use of a well known
fact that organisms seldom respond to these or other climatic factors
singly, but that tolerance to one is usually in some way related to
the value of the other. This can be seen in the accompanying graph
of sheridanii localities ( fig. 2 ) , plotted for values of mean January
and mean July temperatures. When mean July temperature is below,
for example, 60^ F, then sheridanii will occur in such localities only
if mean January temperature is somewhere between, roughly, 14° -
22°; but when mean July temperature exceeds 70° it occurs in
localities where the mean January temperature is above about 24°.
Thus for any given July temperature the range of tolerance to January
temperatures is much less than the range applicable to the species
over its whole range and conversely.
mean Jan.
2(2):in-]60, 196}
CALLOPHRYS FROM NORTHWEST
159
Fig. 2. Graph of known localities of C. {Callophrys ) sheridanii (numbered
as in table 1 ) according to their mean January and mean July temperatures.
160
CLENCH
/. Res. Lepid.
Similar plots (mean January against mean July) have been con-
structed for several other Norther American lycaenids {Callophrys {In-
cisalia) ] Lycaena thoe\ all of which show patterns generally similar
to that of sheridanii. In all, for example, the correlation is rectilinear,
occupying a band of more or less uniform breadth, though this
breadth, the slope of the band, and its position vary from one species
to another. In those species with a sufficient number of locality points
( sheridanii is not one of them ) a further, rather paradoxical, effect is
noted: the upper limit of temperature tolerance is set by a January
threshold, the lower limit by a July threshold. In other words, such
species can extend into warm areas only to a point where increasing
winter temperatures form a barrier; and can extend into cold
areas only so far as summer temperatures continue to be sufficiently
warm. Neither the heat of summer nor the cold of winter seems
to be relevant. This may well be true of sheridanii but the available
data are insufficient to demonstrate it.
There will be, then, within the limits set by the particular factors
singly^ additional "excluded areas’’ in this two dimensional manifold
whose July and January means lie outside this band of correlated
temperature responses. These, when added to the other excluded areas
( as shown on the map, fig. 1 ) , would still further restrict the territory
available to the species. This has not been done for sheridanii because
the number of locality points available is not sufficient to permit a
reliable determination either of the central regression line or the
breadth of the band. Inspection, however, shows that much of the
above-mentioned areas of Montana and South Dakota would thereby
be excluded. This, parenthetically, suggests that the correlated re-
sponse to summer and winter mean temperatures is a major impedi-
ment to the eastward spread of sheridanii.
REFERENCES
BARNES, WILLIAM and FOSTER H. BENJAMIN, 1923. Notes and new
species. Contrib. Nat. Hist. Lejid. N. America^ : 62-96 [Callophrys,
pp. 64-69].
BLACKMORE, E. H. 1927. Check list of the Macrolepidoptera of British
Columbia, Victoria-. E. H. Ban field, 47 pp.
BOWMAN, KENNETH 1919. Annotated check-list of the Macrolepidoptera
of Alberta. Red Deer: Alberta Nat. Hist. Soc., 16 pp.
CLENCH, HARRY K. 1944. Notes on lycaenid butterflies, a. The genus
Callophrys in North America. Bull. Mus. Comp. Zool. 94: 217-229.
1961. Tribe Theclini in Paul R. and Anne H. Ehrlich, How
to know the butterflies (Dubuque, Iowa: W. C. Brown Co.): 177-220
[Callophrys {Callophrys), p.p. 209-210],
LEIGHTON, BEN V. 1946. The butterflies of Washington, Dniv. Washington
Publ. Biol. 9: 47-63.
Journal of Research on the Lepidoptera
2(2) :161-169, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 196}
EUPHYES DUKESI
A Review of Knowledge of its Distribution in
Time and Space and its Habitat
BRYANT MATHER
Box 2131 Jackson, Mississippi
Euphyes DUKESI Lindsey/ has been reported from Alabama,
Virginia^ Ohio, Michigan, and Louisiana. This paper reports its occur-
rence in Mississippi and North Carolina and in a new section of Louisi-
ana, gives notes on its habitat' and on its known distribution in time
and space.
The finding of E. dukesi in Mississippi suggested the desirability
of reviewing available information on all the known occurrences. In
the course of the review, valuable unpublished information on its occur-
rence in North Carolina, Virginia, Louisiana, Michigan, and Ohio was
obtained and is here presented. It is hoped that this review will stimu-
late search for additional occurrences, especially in the wide gaps that
separate the known occurrences.
ALABAMA
Euphyes dukesi was described by Lindsey (1923) from a type series
of four specimens from Mobile Co., Alabama, that were taken by W. C.
Dukes of Mobile. Bell (1926) gave additional data on the type locality
and reported having taken specimens on Hibiscus flowers one mile
from Union Station, Mobile, in a marsh with waist- to shoulder-high
grass. Lindsey, Bell, and Williams ( 1931) reported that it was collected
in considerable numbers in 1925 by Dr. Van Aller and Mr. Lading
(sic)^ of Mobile and by Bell during a short collecting trip near that
city. It was on the wing from 24 August to 11 October. Holland
(1931) figured a S, referred to in his text (p. 388) as "type” and
on his plate (plate LIV, fig. 26) as "paratype” and stated: "It occurs
in southern Alabama and probably elsewhere along the Gulf.” Klots
(1951) figured a pair from Mobile, Ala. Evans (1955) recorded that
2 S $ and 1 $ from Alabama were in the British Museum. Clench
(in litt) reported that the only two specimens of dukesi in the Car-
negie Museum are the two para types mentioned by Lindsey (1923)
as being in his collection.
'This species was formerly placed in the genus Atrytone Scud. 1872. Evans (195 5 noted that
the type species of Atrytone is arogos B. & LeC. 1833. He included Atrytone only arogos and
logan. He also noted that the type species of Euphyes Scud. 1872 is vestris Bdv. He included in
Euphyes the remaining species of North American skippers formerly lumped with arogos and logan
under Atrytone. The validity of the separation of the species formerly so lumped into the two
groups into which Etans divided them, and hence the validity of the change of the generic name
for dukesi from Atrytone to Euphyes appears to be concurred in by all of those with whom the
author corresponded in collecting the information summarized in this paper.
2Henry Peter Loding, author of Catalogue of Beetles of Alabama. Ala. Geol. Surv., Monograph 11,
1945, 172 pp.
161
162
MATHER
/. Res. Lepid.
VIRGINIA
Clark (1932) did not list duke si as possible for the vicinity of
the District of Columbia. Clark and Clark (1951) described the
area in Norfolk and Princess Anne Counties from which they had
dukesi records based on their own work and that of Otto Buchholz.
They concluded that, in this area, dukesi has one brood that flies
between 12 June and 16 July. Martin and Truxal ( 1955) reported
that the Los Angeles County Museum collection included dukesi taken
in Virginia in August, Rindge checked the American Museum of
Natural History series and reported (in Litt) that there were 35 S $
and 26 $ $ . Virginia specimens from the Buchholz collection include
both S S and $ $ taken in June for which the localities are given
as "Norfolk Co.,” "Pr. Anne Co.,” and "Suffolk.” Suffolk, Va., is in
Nansemond Co. A $ , probably collected by Buchholz, that came to
us from C. F, dos Passos, is labelled "Pecaty Road, vie. Hickory, Va.
Norfolk Co., 26 VI 40” Charles V. Covell, Jr. reported (in litt) that
he had not taken dukesi in his collecting in the Norfolk area. Col. S. S.
Nicolay reported (in litt) that he had taken specimens in Virginia in
all months, May through August, is convinced that, in this area, dukesi
is double brooded, and estimated that the first brood is out between 25
May and 5 July and the second between 20 August and 15 September.
He thought that it may be possible for it to be found as late as October.
Col. Nicolay provided the following notes on collecting dukesi in
Virginia: "A fair number of specimens were taken in the period
4-15 July 1951. The location of the habitat is the difficult factor;
once the habitat is located, they are not particularly difficult to
capture. North Landing, Princess Anne Co., a locality mentioned by
Clark and Clark (1951), is actually a bridge crossing a canal. Dukesi
is found abundantly in the few yards of marsh on either side of the
bridge and along the banks of a small stream that empties into the
canal at the bridge, but it is not found 100 yards away from the
bridge on either side, nor in any of the fields a block away. Dukesi was
also found in other spots in and around the Dismal Swamp area of
Princess Anne and Norfolk Counties. Each locality is actually a
Spot since the insects never wander away from their chosen ground.
One can collect in a field or along a trail within a stone’s throw of
the deep swamp habitat of dukesi and not find a single stray. They
were not found more than a few yards from standing water. The
best collecting method is to work the edges of small roads that cut
through the gum swamps. The ^ in search "of 9 9 will fly along
the edge of the road for a few yards before turning- back into the
tall swamp grass from whence they came. This grass grows under the
trees of the swamp, not out in the open. They are fond of pickerel
2(2):161-169, 1963
EUPHYES DUKESI
163
weed {Pondeteria cor data) and at times were found swarming around
clumps of its spired blue flower along the edges of small bodies
of water. The specimens taken in July 1951 were quite well worn,
especially the $ $ , with an occasional $ appearing fresh. On 25
August 1951, at the same localities, freshly emerged specimens were
found in large numbers, particularly $ S . These relations have
been found to exist in each of the five seasons during which obser-
vations have been made.”
OHIO
References to the occurrence of dukesi in Ohio were found in
Freeman (1942), Harris (1950), Klots (1951), and Pliske (1958
”1957”), hut no detailed published account has been located. The
earliest record of dukesi from Ohio that was found is of a ^ taken
on 10 July 1940, at Sylvania by Donald Eff given in Freeman ( 1942) ,
Sylvania is 11 miles northwest of Toledo, in Lucas Co., close to the
Michigan border. Eff described (in litt) the locality as "in the Oak
Openings in northwest Ohio, an acid, quicksand base, area of about 140
square miles set in the midst of rich farming land. Before canals
were dug and the water level was higher, it was a combination of
swamps and higher sandy knolls. Much of the land in the northeast
corner of Indiana and southeast Michigan would fall in the same general
category.” The occurrence of dukesi in Ohio was apparently not
known to Macy and Shepard (1941). E. S. Badger of Kokomo,
Ind., described (in litt) his experiences collecting dukesi in Ohio on
7 July 1957 as "one of the most fantastic experiences I have had in
this section of the country. Homer Price took me to the old disused
bed of the Miami Canal on a blistering hot sticky day. Equipped with
knee-high rubber boots, we followed the canal to an area where it
was over-grown with wide-bladed sedges from bank to bank. Sloshing
around in water 6 to 12 inches deep, we started seeing c? flying
all over the place. The canal was pretty well shaded by trees and
was only about 12 to 15 feet wide. There were some sedges in the
adjoining woods in one area and here, particularly, dukesi abounded.
It had been our intention to take only a half dozen specimens but
the skipper was so abundant that I ended up with 36 $ $ and 1 $
in about 90 minutes of collecting. The only other butterflies were
common dark skippers, a few worn Strymon falacer, and worn Speyeria
cybele.” One $ collected by Badger on 7 July 1957, now in our
collection, is labelled: 'Miami Canal, Paulding Co., Ohio.” Mr. Badger
not only sent us the foregoing account but also asked Mr. Homer F.
Price of Payne, Ohio to provide additional information. Mr. Price gave
(in litt) the following records and comments:
(a) Marie DeLarme Creek, Carryall Township, Paulding Co.,
164
MATHER
}. Res. Lcphl.
Ohio: 29 June 1955 9 , 4 July 1955 3 6 6 , 30 June 1956 6 , 8
July 1956 6 , the sedges are nearly shoulder-high some years at this
station, low ground along the creek. A small colony; in some seasons
none are seen.
(b) Miami Canal, Defiance Township, Defiance Co., Ohio: 8
July 1956 S . Very wet woods along the canal, the sedges were
rather short, none were seen in the canal, which was dry.
(c) Miami Canal at Charloe, Brown Township, Paulding Co.,
Ohio: 14 July 1950 c? 9,16 July 1950 2 o o , 23 July 1950 2 5 c?
30 July 1950 9, 25 August 1950 5 (worn, latest record), 2 July
1959 3 5 5 19 (fresh). 'The species is quite common some seasons
in and along the canal where the sedges are of normal size. They
occur at rather open spots where there is a little sunshine. Many
specimens have been collected at this station.
"I am of the opinion that this species is more plentiful in this
area than is generally believed. Recently I have collected dukesi at
various stations along the canal from the Defiance County line .south
to the Putnam County line. This area has a low gradient, having been
a part of the Black Swamp. The soil is a very heavy clay. I have been
of the opinion that dukesi probably appeared first in v/estern Ohio
near the southern terminus of the Miami Canal at Cincinnati and
extended its range northward along the canal. Hot weather, mud
and water, mosquitoes and deer flies, probably cause some collectors
to remain at home during early July when dukesi should be searched
for. It appears that we have only one brood here. They seem to
like shade - but not deep shade.”
Badger gave (in litt) the following additional Charloe records
from specimens now in his collection that were taken by Price: 15
July 1951 2 5 5 , 22 July 1951 9, 21 July 1954 9, and 4 July
1955 5 . Badger has included dukesi on his preliminary manuscript
list of Indiana butterflies on the basis of its known occurrence in Ohio
within a few miles of the Indiana border. Price also mentioned that
he had correspondence with Chermock and had provided material
for study in connection with the possibility that the Ohio and Alabama
populations might be subspecifically different. Coveil stated (in litt)
that his collection contained a 5 obtained from Frank Chermock
labelled "Chasloe, Pauling Co., Ohio, 15 July 1957” taken by Price.
LOUISIANA
Lambremont (1954) described the single speciment known to him
from Louisiana, taken by him, as constituting a possible westward
range extension. His specimen^ a 5 , Tulane University #2064, was
taken IIV2 miles west of Oak Grove, West Carroll Parish, on 19
June 1950. This locality is near the Boeuf River about 20 miles west
2(2);161-169, 196}
EUPHYES DUKESI
165
of the Mississippi River, and more than 200 miles north of the Gulf
of Mexico. Harris (1950) gave the range of dukesi as: "known
from southern Louisiana, southern Alabama (Mobile), soudieastern
Virginia, and Ohio.’’ Harris wrote (in litt) that the information on
dukesi given in his 1950 report was v/ritten by A. H. Clark and has
kindly provided a copy of Clark’s letter of 13 July 1950 to him,
reading in part; "I am returning the manuscript ... I have added
. . . in brackets ... A. dttkesi which certainly occur (s) in Georgia.’’
Harris commented, with regard to Clark’s reference to "southern
Louisiana” that he assumed that Clark knew of records. It seems un-
likely that Clark on 13 July 1950 would have known of Lambremont’s
19 June 1950 record. .The reference by Clark is almost certainly to
a previously unpublished occurrence discovered by William D. Field
which Mr. Field has described (in litt) as follows: "I took a series of
dukesi near a drainage ditch in what had once been a swamp near
Camp Plauche, Jefferson Parish, La., on May 15, 1944. This swamp
had been well-drained but had a good flora resembling a woods with
only a light amount of undergrowth. The details of the latter escape
me except for the catbrier {Smilax). Camp Plauche was right next to
the town of Harahan, less than a mile from the Mississippi River. This
series was sent to Austin Clark for the museum. However, at present
there is only a single male in the collection.” Harahan is on the
north (left) bank just a little upstream from New Orleans.
MICHIGAN
Pliske (1958 "1957”) described the occurrence of dukesi at Ann
Arbor, Michigan. He reported 2 S S taken by Arthur Slater and
himself on 21 July 1956 in a small marsh on the north side of the
Huron River about one mile east of Ann Arbor. Washtenaw Co.,
Mich.; a second $ taken at the same locality the following day; and
a $ taken on 13 July 1957. A second locality, Vj mile east of the
first, yielded a 2 on 27 July 1956 and a $ on 15 July 1957. Both
marshes were dominated by the lake-margin sedge {Carex lacustris)
characterized by broad blades, 1 cm. or more in width, spreading in
a criss-cross manner. On 28 July a 2 was observed ovipositing on
the underside of a sedge leaf about HA feet above the ground. The
eggs were collected by W. H. Wagner and were described by Pliske.
Pliske’s account refers to the Ann Arbor colony as the fourth to have
been discovered, actually, although not known to him, it was the
sixth, the fourth and fifth having been the two Louisiana occurrences.
Wagner has provided (in litt) additional data on the Michigan occur-
rence of dukesi as follows: Subsequent to the development of the
data reported by Pliske, a battered 6 that had been taken by Sherman
Moore on 28 July 1952 at Highland Park, Oakland Co., Michigan
166
MATHER
/. Res. T.epitf.
was found in the University of Michigan collection in the E dion
series. Wagner notes that the thoroughly urbanized region of Highland
Park in Detroit is wholly unsuited to the occurrence of dukesi and
that the specimen must be a stray. Wagner having visited the North
Landing, Virginia, locality with Clark and also the Ann Arbor, Mich-
igan locality, compared them as follows; 'The habitat there (North
Landing) was very different from the Michigan one: deep, shady
swamp, the butterflies rather slow flying among grasses and sedges.
Here the species reminds me more of dion. It flies with the latter
as well as A. logan, P. viator, E. conspicua, and P. massasoit!’
MISSISSIPPI
Mather and Mather ( 1958) regarded L. dukesi as of probable
occurrence in Mississippi. This was proved correct when I took a $
on 4 October 1959 about 4 miles north of Clinton, Hinds Co. The
locality is on the north side of the Kickapoo Road bridge over Bogue
Chitto Creek, Sec. 3.5, T 7 N, R 1 W. The specimen was taken at
about 3 p.m., feeding on flowers of Eupatorium {Conodinum) coel-
estinum that were growing in a marshy area within a few feet of a
pool of essentially stagnant water in the creek bed. This locality is
about 200 miles north of the Gulf of Mexico. A comparison of the
three males at hand reveals that the Mississippi specimen is more like
the Virginia one than the Ohio one especially in that it has less yellow
hair on the hindwing above and less bright yellow on the hindwing
below. The separation of the black areas of the stigma on the fore-
wing above is most distinct in the Virginia specimen, slightly less
distinct in the Ohio specimen, and still less distinct in the Mississippi
specimen. At the same locality on the same date, the first Mississippi
specimen of Poanes viator was taken. A second viator was taken there
on 11 October 1959 by M. & E. Roshore but no additional specimens of
either species have been found on numerous subsequent visits.
NORTH CAROLINA
Nicolay reported (in litt) the taking of dukesi on 6 June I960
at a locality about 7 miles southeast of New Bern, Craven Co., in
a small swamp located along the Neuse River. Based on his observa-
tions in Virginia he regarded the locality as a typical dukesi habitat.
Specimens of both sexes were taken, the condition of the $ $ sug-
gested that they had been on the wing for about a week, the $ $
were quite fresh.
2(2)J6l-169, 196}
EUPHYES DUKESI
167
SUMMARY AND DISCUSSION
The geographical distribution of the localities at which E. dukesi
has been found to occur is shown by figure 1; also shown are the wide
areas between the Great Lakes, the Middle Atlantic Coast, and the Gulf
States from which the insect is not yet known. Nicolay reported (in
litt) that he did not find dukesi during the two years (1 947-49 )
that he collected in the Pensacola, Fla., area. Zeiger reported (in
litt) the taking of a series of 17 c? ^ and 5 9 $ of P. viator on
23 May 1962 among giant cutgrass, in brackish water, along the edge
of the Suwanee River, near its mouth. He will visit this locality
again and check for the occurrence of P. dukesi. Neither Lambremont
(1954) nor Mather and Mather (1958) found it in coastal Louisiana
or Mississippi. Freeman (1951) did not report it from Texas. Harris
reported (in litt) that he felt certain that it will be found in swamps
in Georgia such as those of the Altamaha River, the Okefenokee,
and other suitable localities. He also indicated that Buchholz, whose
records of dukesi from Virginia were cited by Clark and Clark (1951)
and who had many Georgia records, apparently had no records of
dukesi from Georgia. Roever (in litt) reported that he had never
collected in the type of marsh in Tennessee in which dukesi might
be expected to occur but regarded the open marsh around Reelfoot
Lake in northwest Tennessee and the spring-fed gum swamps in
the Hatchie River bottom near Viedo, 4-5 miles southeast of Mercer,
Madison Co., as likely areas.
E. dukesi appears to be single brooded in Ohio and Michigan and
double-brooded in Virginia and the Gulf states; the months of record
for the seven states in which it is known to occur are shown in table 1.
E dukesi appears to be a swamp butterfly, but while the Ohio
and Michigan occurrences are apparently closely connected with sedges,
the Virginia occurrence is in the gum (Nyssa aquatica) swamps, which
as Clark and Clark (1951, p. 8) pointed out, lie west of the sedge
marshes that, in turn lie behind the sand dunes of the Virginia coast.
Until more complete knowledge of the distribution of dukesi becomes
available, it would seem prudent to describe its distribution as: north-
east and southeast Louisiana, central Mississippi, southwestern Ala-
bama; eastern North Carolina, southeastern Virginia; northwestern
Ohio, and southeastern Michigan, rather than in terms that imply its
occurrence in the large regions between these areas of known occur-
rence.
168
MATHER
J. Res. LepiiJ.
Fig. 1. Map showing the locations where Euphyes dukesi
has been found.
MAY
JUN
JUL
AUG
SEP
OCT
MICHIGAN
X
OHIO
X
X
X
VIRGINIA
X
X
X
X
(a)
(b)
NORTH CAROLINA
X
ALABAMA
X
X
X
MISSISSIPPI
X
LOUISIANA
X
X
(a) Probable (b) Possible
TABLE 1. The monthly occurrence of Euphyes dukesi at
various localities in the eastern United States.
2(2): 161-169, I96.i
EUPHYES DUKESI
169
ACKNOWLEDGEMENTS
The principal value of this paper derives from the data provided
by Col. S. S. Nicolay, Cherry Point, N. C.; Mr. William D. Field,
Washington, D. C.; Mr. F. S. Badger, Kokomo, Ind.; Mr. Homer F.
Price, Payne, Ohio; Mr. Lucien Harris, Jr., Atlanta, Ga.; Mr. Charles
V. Coveil, Jr., Norfolk, Va.; Dr. Frederick H. Rindge, New York,
N. Y.; Mr. Charles F. Zeiger, Jacksonville, Florida; Mr. Harry K.
Clench, Pittsburg, Penn.; Mr. Kilian Roever, Tucson, Ariz.; Dr.
Warren H. Wagner, Ann Arbor, Mich; Mr. Donald Eff, Boulder,
Colo.; Mr. H. A. Freeman, Garland, Texas; Mr. John M. Burns, Middle-
town, Conn.; and their kind permission to publish them; their coopera-
tion is deeply appreciated.
REFERENCES
BELL, E. L. 1926. Notes of some Hesperiidae from Alabama, Jour. N. Y.
EnL Soc., 34: 269-271.
CLARK, AUSTIN H. 1932. The Butterflies of the District of Columbia and
vicinity. Smithson. Inst., Bull. U. S. Nat. Mus., 157, 337 pp.
CLARK, AUSTIN H. and LEILA F. CLARK 1951. The butterflies of Vir-
ginia. Smithson. Misc. Coll., 116 (7): 1-239.
EVANS, W. H. 1955. A Catalogue of the American Hesperiidae, Part IV.
British Museum- (London) , 499 pp.
FREEMAN, H. A. 1942. Notes on some North American Hesperiidae with
the description of a new race of Polites verna ( Edwards ) ( Lepidoptera ;
(Rhopalocera) . Ent. News, 53: 103-106.
_ — 1951. Ecological and sytsematic study of the Hesperiidea or
Texas. Sotithern Methodist Univ. Studies, No. 6: 1-67.
HARRIS, LUCIEN JR. 1950. The butterflies of Georgia. Bull. Georgia Soc.
Naturalists, 5 : 29 pp.
HOLLAND, WILLIAM J. 1931. The Butterfly Book (revised edition).
Douhleday & Co., Garden City, N. Y., 424 pp.
KLOTS, ALEXANDER B. 1951. A Field Guide to the Butterflies. Houghton
Mifflin Co., Boston, 349 pp.
LAMBREMONT, EDWARD N. 1954. The butterflies and skippers of Louis-
iana. Tulane Stud. ZooL, 1: 125-164.
LINDSEY, A. W. 1923. New North American Hesperiidae. Ent. News.
34: 209-210.
ERNEST L. BELL and R. C. WILLIAMS, JR. 1931. The Hes-
perioidea of North America. Jour. Sci. Lab. Denison Univ. 26: 1-142.
MACY, RALPH W. and HAROLD H. SHEPARD 1941. Butterflies. Univ. of
Minnesota Press, Minneapolis, 247 pp.
MARTIN, LLOYD M. and FRED S. TRUXAL 1955. A list of North Amer-
ican Lepidoptera in the Los Angeles County Museum, Part 1 - Butterflies.
Los Angeles Museum, Science Series, No. 18: 1-35.
MATHER, BRYANT and KATHARINE MATHER 1958. The butterflies of
Mississippi. Tulane Stud. Zool. 6: 63-109.
PLISKE, THOMAS E. 1958 ”1957”. Notes on Atrytone dukesii (sic), a rare
species new to southern Michigan (Hesperiidae). Lepid. News, 11: 42.
Journal of Research on the Lepidoptera
2 (2) :170-172, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 1^6}
THE COMPLETE LIFE HISTORY OF STAPHYLUS
HAYHURSTI
JOHN RICHARD HEITZMAN
3112 Harris Ave., Independence, Missouri
This obscure little skipper is one of those species that are
easily overlooked by the non-observant collector. It inhabits a wide
range in the eastern United States extending at least as far as Pennsyl-
vania and Nebraska and south to Florida and Texas, being replaced
in southern Texas by the closely related Staphylus mazans Reakirt. Host
plants include Chenop odium and Alternanthera and the species is rather
localized to areas where these weeds grow. One of the peculiarities of
hayhursti is the moth like habit of spreading the wings out flat against
the surface upon which it alights. It is strongly attracted to certain
flowers and at times I have found a dozen or more specimens clustered
about a bed of Spearmint flowers {Mentha spicata) during late July
in a good local. Sweet Clover (Melilotus) also seems to have a par-
ticular attraction for the species. In rural areas the best collecting
spots for hayhursti will be associated with moist rather shady habitat.
An idea spot is a small creek running through a wooded area or the
weedy edges beside an old railroad track. In urban areas it is drawn
to vacant lots grown up in weeds where an ample supply of the host
plants can be found growing. There are two distinct broods produced
each year in Missouri and probably at least three further south. In a
normal year you can find hayhursti from the third week in May to
the end of the third week in June and again from mid July until the
end of August. The larvae produced by the second brood normally
hibernate in the third instar in this area. An occasional third brood
specimen is produced but these are very rare.
From time to time scattered bits of information on the early stages
of this species have appeared but as far as I can ascertain there has
been no complete life history published on this butterfly. In May
of 1962 I was able to obtain 49 ova from a worn female placed on
potted Lambs Quarters and covered with a bag of Nylon netting. From
these eggs 21 adults were reared and the larvae were carefully observed
and recorded in detail. Since Edwards’ description of hayhursti in 1870
was based on material from 'Missouri” this description should approx-
imate topotypical specimens. The illustrations which accompany this
description were drawn by William H. Howe.
170
2(2:)170-172, 196}
STAPHYLUS HAYHURSTI
171
OVA: Eggs were obtained from 25 to 28 May, The eggs were
laid singly usually beneath the leaves with a few placed above and a
few others on the stems. The ova is very small, less than 1 mm. in
diameter and has a flattened circular appearance. The color is a deep
orange brown with a raised cream colored pattern of irregular ridges.
The six largest radiating out from the top center to the sides. Under
low power magnifaction the egg has the appearance of a white star
on an orange background.
FIRST INSTAR LARVA: The first emergence occurred on 1
June. The newly hatched larva is translucent light orange, abdominal
segments 8, 9 and 10 are dull brown dorsally. The head and pro-
thoracic shield are shiny black. After the first day of feeding the body
assumes a pale green color with orange edges. The first meal is a hole
eaten through the top of a leaf. The second day a small tent about
one eighth inch square is made near the tip of a leaf by cutting three
sides and folding the severed section to the top of the leaf and fast-
ening it with a few threads of silk.
SECOND INSTAR LARVA: Body dark green, abdominal seg-
ments 9 and 10 pale orange brown, abdomen translucent orange yel-
low. Head and prothoracic shield clear black. The larvae leave the
tent a short distance and feed during the day on top of the leaves,
eating holes through the leaves. The larvae are very agile and when
dismrbed they rapidly retreat to the tent.
THIRD INSTAR LARVA: Body pale watery green with no vis-
ible markings, thickly covered with short white setae, a few longer
hairs on the last segment. Abdominal segments 8, 9 and 10 are creamy
yellow with a pale brown tone dorsally. The neck is a slightly paler
green than the body. Due to the translucence of the body the edges
of the abdomen appear to be pale yellow. The prothoracic shield has
a pale green dorsal area shading into deep purple at the sides. Head
deep purple and thickly covered with short white setae, face deeply
cleft vertically at the epicranial suture. Mandibles are a contrasting
shade of reddish purple.
FOURTH INSTAR LARVA: Body medium green covered with
short white setae^ abdominal segments 8, 9 and 10 slightly paler, section
10 with a few longer hairs. The neck is slightly paler and the pro-
thoracic shield is visible only as two pale brown subdorsal spots. The
head is deep purple, almost black and thickly covered with medium
length white hair. The growth rate is greatly accelerated in this instar.
FINAL INSTAR LARVA: Length of fully grown larvae, 20-23
mm. Body ground color is a deep green with a rosy overcast, inter-
segmental folds pinkish green. Legs cream color with the rest of the
abdomen and the prolegs deep green. The body is thickly covered with
fine white hair, some slightly longer on the last segment. The spiracles
are pale creamy white and slightly raised. The skin is quite trans-
lucent, the heart visible as a deep gr^en mid dorsal line. In this instar
172
HEITZMAN
J. Res. Lepid.
the prothoracic shield has turned to a very pale brown color. Head
deep purple almost black, thickly covered with white hair and deeply
cleft vertically at the epicranial suture. The tent in the final instar
consists of an entire leaf folded together and pupation occurs in the
tent with the addition of a very thin lining of silk and a silken pad
to which the cremaster is attached.
PUPA: First pupation occurred 27 June. Length of pupae 13-17
mm., width at widest point of wing cases 3-5 mm. The pupa case is
thickly dusted with a white powdery bloom. Abdomen pale orange
brown, the wing cases and thorax light olive brown. Spiracles dark
brown and noticeably raised. The abdomen and head are thickly covered
with short orange bristles. Cremaster dark brown consisting of a single
sharp point. Adults emerged from the 10th to l6th of July, males and
females emerging at the same time. All specimens were quite large
and the measurements for the larvae and pupae given should compare
closely with those of wild specimens. The time spent in each instar
was quite constant, usually five days elapsing between moults.
Fig. 1. Early stages of Staphylus hayhursti. Ova, pupa, mature larva
and enlarged front view of the head.
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Vohime 2
Number 2
September, 1963
IN THIS ISSUE
Quantitative analysis of certain wing and genitalia characters
of Pieris in western North America
Vincent C.S. Chang 97
Biographical sketch — Vincent C.S. Chang 126
Genetic and environmental variation in Pieris brassicae
Brian O.C. Gardiner 127
Type localities of the Megathymidae H. A. Freeman 137
Biographical sketch — H. A. Freeman 142
The distribution of an endemic butterfly Lycaena hermes
Fred Thorne 143
Callophrys (Lycaenidae) from the Northwest
Harry K. Clench 151
Euphyes dukesi Bryant Mather l6l
The complete life history of Staphylus hayhursti
John Richard Heitzman 170
THE JOURHAL
ONJ THE LEFIJPOFTER^A
Volume 2
November, 1963
Number 3
THl J)@UR!NJA\L ©F FISIAK’GH OIMJ
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Journal of Research on the Lepidoptera
2(3) :173-184, 1963
1140 W. Orange Grove Air., Arcadia, California, U.S.A.
© Copyright 1961
AFFINITIES AND DISTRIBUTION OF
ANTILLEAN ITHOMIIDAE’
RICHARD M. FOX
Carnegie Museum, Pittsburgh, Pa.
The family Ithomiidae, a characteristic component of the lepi-
dopterous fauna of the American humid tropics, is represented in the
Antilles only by two species, both members of the genus Hymenitis.
One species is found on Cuba, the other occurs on Jamaica and Hispan-
ola (fig. 1), and their affinities are with Central American species It
is surprising that no other species have reached the islands since the
lands surrounding the Caribbean support a rich ithomine fauna: it is
puzzling to find Hymenitis, a genus of frail sun-shunning species, on
the Greater Antilles instead of any of the sturdy sun-loving genera.
That no ithomine is found on any of the Lesser Antilles is probably
because no precisely suitable ecologies are available on those islands.
In order to explore the zoogeographic implications of the Antillean
ithomines, it is first necessary to review the systematics and phylogeny
of these insects.
SYSTEMATICS
\Hymenitis Anonymous, 1807; column 1180. Type of genus, Papilio
polymnia Linne, 1758, by designation of Hemming, 1934]
Hymenitis Hiibner, 1816: 8. Type of genus, Hymenitis diaphane:
Hiibner, 1816, designated by Scudder, 1875.
—Greta Hemming, 1934: 28. Type of genus, Hymenitis diaphane:
Hiibner, by original designation.
Hemming’s discovery (1934) that Hymenitis first appeared in
print in 1807, nine years prior to the "Verzeichniss,” would seem to
reduce the Hiibner name to homonymy. By selecting Papilio polymnia
as type of Hymenitis Anonymous, Hemming made the name an abso-
lute synonym for Mechanitis Fabricius, 1807; Greta was proposed to
replace Hymenitis Hiibner. Hemming’s actions were justified under
a strict interpretation of the International Code, which provides for
the invalidity of anonymous publications only if issued after 1951.
The 1807 article was an unsigned review of a manuscript by
Hiibner circulated for the information of his colleagues. The anonyrcious
’This study was conducted with the support of National Science Foundation
grant GB-510.
173
174
FOX
/. Res. Lepid.
a
Antillean ithomines and related species.
2(}):I73-1S4, 1963
ANTILLEAN ITHOMIIDAE
175
reviewer saw fit to publish the contents of the Hiibner manuscript, but
with some unauthorized alterations: some of the proposed generic
names were replaced, others were juggled. One of the names juggled
was Hymenitis and the reviewer used it for insects other than those
Hiibner intended.
The two objections to accepting the validity of this publication are
that the generic name Hymenitis has no author, and that the unsigned
article was malicious and "scooped” Hiibner in a manner which today
would be condemned as being unethical.
Hemming avers that the review was in fact written by Illiger, the
editor of the journal, but there is no proof of authorship in the article
itself. At most, Illiger as editor was responsible for permitting an
unsigned review to appear, but responsibility and authorship are not
the same things. Hymenitis Anonymous was the illegitimate offspring
of a mother (the editor), but no father (author) was willing to claim
it. Proving that Illiger was the author of the review would not purge
it either of anonymity or of unethical intent. In my opinion, the first
legitimate publication of Hymenitis as a generic name was by Hiibner,
1816. Hemming could have cleared the situation best by requesting
that the 1807 article be suppressed.
HYMENITIS CUBANA
Hymenitis cubana Herrich-Schaf fer, 1 862 : 1 1 8. ( Cuba ) .
Figs. 2 and 7
Gundlach (1881) recorded the larval food plant as galan (the
local name for a member of the genus Ce strum, family Solanaceae)
and observed that the insect flies slowly and is nearly invisible in the
forest shadows. He reported cubana from the mountains at each end
of the island and said it seemed to be absent from the central Trinidad
range; it since has been captured in the central mountains.
Specimens exmined, 6 ^ , 12 $ :
Cuba: Sierra Maestra, Oriente, 1000 feet, 1^,29 (M.C.Z. )^
Loma del Gato, Cobra Range, Oriente, 3000 feet, 2 , 4 9 (M.C.Z.).
Turquino River, Oriente, 1 9 (M.C.Z.). Buenos Aires, Trinidad Mts.,
2500-3500 feet, 1 ^ (M.C.Z.). No station (Oriente, Bates, 1935 ) ,
2^,59 (M.CZ.).
HYMENITIS DIAPHANA
Heretofore known only from Jamaica, a second subspecies from
Hispanola is described below. H. diaphana is easily distinguished from
H. cubana because in the former the black marginal band of the fore-
wing follows the wing contour at the apex, while in the latter the black
margin is wider and heavier at the apex in males and in females fills
^The following abbreviations are used to indicate museum collections :
A.M.N.H., American Museum of Natural History, New York; C.M., Carnegie
Museum, Pittsburgh; M.C.Z., Museum of Comparative Zoology, Cambridge.
176
FOX
/. Res. Lepid.
the entire area beyond the white postdiscal band. Despite these external
differences (figs. 2 and 3) the male genitalia of the two species are
similar (figs. 7 and 10), though ciiaphanu has the saccus, penis and
uncus shorter and a different armature at the apex of the valve.
Hymenitis diaphana diaphana (Drury), 1773: 13; pi. 7, fig. 3.
( Jamaica ) .
Figs. 3 and 10
Drury’s figure accurately depicts a male and there can be little doubt
of the correct identification. Avinoff and Shoumatoff (1048) comment
that diapha7ia is "extremely localized, restricted sometimes to a stretch
of land a half-mile across. In such places it is very abundant. It never
emerges into the bright sunlight from its dense humid habitat.’’ It is
found only in the Blue Mountains of eastern Jamaica at about 3000
feet above sea level.
Fig. 2. Hymenitis cuhana, a male from the Trinidad range, Cuba.
Fig. 3. H. d. diaphana, a male from Corn Puss Gap, Blue Mountains,
Jamaica.
Fig. 4. H. d. quisqueya new subspecies, holotype male from Mt. Diego
de Ocampo, Dominican Republic.
Fig. 5. H. polissena umhrana, a male from Cachi, near Mt. Irazu, Costa
Rica.
Fig. 6. H oto. a male from Mt. Irazu, Costa Rica.
2(}):l7)-n4, 1961
ANTILLEAN ITHOMIIDAE
177
Specimens examined, 74 6 , 45 9 :
Jamaica: Corn Puss Gap, 37 (5 , 10 9 (C. M., M.CZ. ). Cuna Cuna
Pass, 31 (5 , 34 9 (C.M., M.C.Z.). Fish Brook, 1 (5 , 1 9 (C.M. ).
Moore Town, 2 S (C.M., M.C.Z.). John Crow Hill, 1 6 (C.M.).
No station, 2 c? ( A.M.N.H., M.CZ.) .
Hymenitis diaphana qmsqneya new subspecies
Figs. 4 and 11
Three males taken by Darlington in the mountains of northwestern
Dominican Republic between 3000 and 4000 feet above sea level in
July represent an undescribed subspecies and extend the known range
of the species to one more island of the Greater Antilles. It is possible
that diaphana may eventually be found on Puerto Rico.
The Dominican subspecies is similar to H. d. diapha^ia of Jamaica
but the following differences are noted in males: The dark discocellular
bar of the forewing is narrower and the white costal spot and white
scaling on the base of Mi is only half to two thirds the width of the
same markings in H. d. di'tphana. The black marginal scaling is a little
bit wider between M3 and CUi of the forewing and betv/een M3 and
CU2 of the hindwing, so that this band gives the impression of being
of a more uniform width than is true of the Jamaica subspecies. On
the underside all dark markings are yellowish brown and appear to be
paler than the reddish brown present on H. d. diaphana.
Figs. 7 to 12. Male genitalia in lateral view, the left valve removed
and the dissected penis below. Fig. 7, H. cuhana. Fig. 8, H. oto. Fig 9,
H. morgane. Fig. 10, H. d. diaphana. Fig. 11, H. d. quisqueya new subspecies,
paratype. Fig. 12, H. polissena umbrana.
178
FOX
/. Res. Lepid.
The male genitalia do not differ.
Holotype $ and two $ paratypes: Mt. Diego de Ocampo, Domin-
ican Republic, 3-4000 feet, July 1938, Darlington. Holotype and one
paratype in Museum of Comparative Zoology; one paratype in Car-
negie Museum (male genitalia number 1172J).
The ancient indian name for Hispanola is said to have been Quis-
quey.
PHYTOGENY
The coloring and pattern of the wings are difficult to use to associate
ithomine subspecies correctly (Fox, 1956) and for that reason must be
used with caution to analyze phylogenies. As in many other genera of
this family, the wings of Hymenitis are mostly transparent and the
markings are reduced to the dark marginal bands on both wings, the
discocellular band of the forewing and some white scaling beyond the
forewing discal cell and faint white spots against the marginal bands.
That so many transparent ithomines look so much alike is not part of
the "mimicry” phenomenon: transparency is a protective adaption of
the species that live within the shadows of the dense forests.
Study of the Ithomiidae demands close attention to morphologic
detail both for correct identification and for phylogenetic analysis. The
venation of the hindwings suffices to define most of the genera, but
to evaluate the relationships among species of a genus, the male genitalia
usually are more useful than wing pattern.
Hymenitis diaphana and H. cuhana differ from all other members
of the genus by the extraordinary length of both the penis and the
saccus (figs. 7, 10 and 11). These two species share a simply formed
triangular valve with two other species, both Central American, H. oto
(fig. 8) and H. morgane (fig. 9). All other Hymenitis species have
the sacculus of the valve greatly enlarged and produced posteriorward
as a prominent projection which may be quite complex in some. H.
polissena umhrana Haensch has this development of the sacculus pres-
ent in a less complex form (fig. 12), and in this respect connects the
diaphana group with the rest of the genus.
The pattern of H. cuhana, although so strongly reduced, has one
pecularity of interest: the black marginal band of the forewing is
exceedingly thin in Cui-Cu2, but wide at the anal angle and above
Cui and in females the marginal color fills most of the wing apex be-
yond the white transverse band. This arrangement of the marginal
coloring occurs in only one other species of Hymenitis, Central American
oto. Because of the similarities between cuhana and oto with respect
both to male genitalia and to pattern, it is evident that these two species
are closely related. Because of certain structural features present in
cuhana and not in oto — the length of the penis and saccus in males,
and the unique shape of the forewing, which is produced to angles
at M3 and at the apex — oto must be the more primitive. This by
2(}):I7}-IS4, 196}
ANTILLEAN ITHOMIIDAE
179
no means indicates that cub ana necessarily evolved from oto; it indi-
cates only that the two species were derived from common stock and
that cuhana has become the more specialized, perhaps because its limited
geographic range would favor the establishment and preservation of
mutations better than the wide range of oto.
The residual black markings of diaphana are narrowed to a degree
matched only by polissena of Costa Rica and western Panama {H. p.
umbrana) and western Eucador {H. p. polissena). Superficially the
two species look exactly alike, .but in diaphana the discocellular veins
of the fore wing are displaced distad so that the discal cell is about two
thirds of the wing length, while in polissena and all other Hymenitis
the discal cell is only about one half the wing length. Despite the
almost exact duplication of pattern between diaphana and polissena,
which certainly cannot be mimicry, the male genitalia differ markedly
Not only is the shape of the valve in polissena (fig. 12) more com-
plex, but the penis is of an entirely different shape, being slender,
poorly sclerotized and deeply sinuate. H. polissena and diaphana are
not nearly as closely related to each other as are cub ana and oto. The
male genitalia of morgane ( an exclusively Mexican species ) are similar
to those of diaphana, but the patterns on the wings differ greatly.
ZOOGEOGRAPHIC CONSIDERATIONS
The vertebrate fauna of the Greater Antilles seems in general to be
related to and derived from the Central American fauna, according
to Darlington (1957) and Brown (study in preparation) finds that
the butterfly fauna of Jamaica is similarly related; the Antillean Itho-
miidae fall into the same pattern. The zoogeographic problem is to
ascertain how the fauna of the Antilles got there from Central America
and to explain why the South American species are so poorly repre-
sented.
For many years it was believed that the Greater Antilles had been
physically continuous with Central America until well into Tertiary
time and that the existing Antillean fauna was derived, at least in
part, from relicts marooned on what became islands when the seas
intruded. Schuchert (1935) held this view.
It is no longer believed that when a faunal relationship is shown
between two areas separated by seas, such a relationship automatically
demonstrates the former existence of a land connection. Simpson
(1940, 1953), Darlington (1957) and others have demolished most
of the bridges erected by earlier zoogeographers, and only a few such
connections are now acceptable. Speculation involving continents drift-
ing about has been put forward in various forms, but continental drift
has been used chiefly as a plausible ( ) substitute for intercontinental
land bridges which otherwise would have had to be discarded.
180
FOX
J. Res. LepiJ.
Animal distribution is generally explained without resort to "Lost
Atlantis” or to continental drift. Matthew’s theory (1915) of hol-
arctic distribution accounts adequately for the presence of living faunae
on all continents and islands; within the framework of Matthew’s
theory, rafting accounts for much of the dispersal to islands. The term
"rafting” may be taken in a general sense, comprising transportation
by literal rafts, by aerial means and by "hitch hiking.”
Rafts of flotsam, formed in flooded rivers and drifting out to sea,
carry many animals. Should such a raft happen to find its way to
an island, the surviving individuals could disembark and establish them-
selves in a new habitat. Transportation by aerial means is well known
for spiders and occurs in many other groups, especially for small, light
weight arthropods. The power of flight combined with air currents
may carry somewhat larger species for great distances. Hitch hiking is
usual for arthropods with parasitic or semiparasitic habits.
The hazards involved in rafting by any means are such that the
chances of a successful journey — one leading to a new colony of the
species — are small indeed, and the hazards are especially great in the
flotsam rafts. Flotsam rafts normally break apart at the mouth of the
river and at sea they are likely to be well doused by salt water even
if no storm is encountered. Should rafted individuals survive exposure
and the vagaries of wind and current happen to put them ashore, the
landing must be made at or very near a suitable ecological situation.
It has been pointed out that, despite the poor probabilities, a succesful
trip need occur but once during a very long period of time and it is
not difficult to demonstrate mathematically that fortuitous if im-
probable conditions might pertain occasionally. If the animal being
rafted is physiologically able to survive exposure to salt water, or if
its habits are such that it may travel in some protected part of the raft,
there is a better chance of a successful journey. Presumably most
mammals, for example, need only to cling to the flotsam until a landing
is made and during the trip they would face a greater hazard from
starvation or exposure to sun than from salt water; the larvae of wood-
boring insects might be transported safely within their burrows. Con-
versely, animals without physiological protection against exposure and
without habits that would place them in a protected situation en route,
have a lessend probability for survival.
Transportation by flotsam rafts appears to be out of the question
for Hymenitis. Salt water would destroy eggs, larvae, pupae or adults
and none of these stages is passed in a protected place. Because
Hymenitis requires a very specific ecological situation, one which is
not present near the Antillean coasts, no mathematical calculation of
the chances of survival for a long enough time to reach an island has
any bearing. If not destroyed by the sea, these insects would still face
an overland journey of many miles in order to reach forests 3000 feet
above sea level. Furthermore, it seems implausible that ocean currents
2(3);173-184, 1963
ANTILLEAN ITHOMIIDAE
181
or the winds (fig. 1) could give much assistance to transporting
Central American butterflies to the Antilles.
Aerial transportation also is unlikely in the case of Hymenius,
though it cannot be ruled out as a mode of dispersal for other butter-
flies. Unlike most groups, HymeniUs remains within the cover of
dense forest and does not seek the sun or open places; the reaction
to the least breeze is to drop to the ground and cling. It is conceivable
that a hurricane could transport adults, provided the wind were strong
enough to suck the insects out of the forest, but not only is the direc-
tion of winds toward, not from Central America, but it is hard to
imagine that such wind force would not also crush and kill the insects.
If rafting by any rhethod could serve to distribute Hymenitis or
the other ithomines, one would expect on the Greater Antilles to find
representatives of the rich fauna of Colombia, Venezuela and the
Guianas rather than of the reduced fauna of Central America, since
all factors of wind and current are favorable to rafting from the
northern coast of South America but unfavorable to rafting from
Central America. Of all the ithomines, Hymenitis and other groups
with similar habits are the most unlikely to be rafted. If rafting
occurred, why are the sun-loving Mechanitis, Melinaea, Tithorea and
Hypothyris not present on on the Antilles?
Darlington (1957) comments, "It might be supposed that the
Gulf Stream would prevent drift from reaching the West Indies from
Central America and might favor drift transport from South America
in spite of the greater distance. But this plausible idea is not supported
by facts.” I must agree with his view on this matter, but for a reason
different than he expressed it. For Ithomiidae there are no facts to
support drift (rafting) by any method.
The Greater Antilles stand within the continental shelf, separated
from Central America only by shallow seas, and the possibility cannot
be dismissed that the two present land areas were once continuous.
Such a connection would not be "Lost Atlantis” bridging the deep
seas and while there is little evidence to support the connection, there
is no geologic evidence against it. Darlington (1957) assumed that
the Greater Antilles were formed by oceanic volcanoes because volcanic
rocks and marine limestones are present. Woodring (1954), however,
commended that, "It is quite evident on ordinary geological grounds
that a considerable part of the Caribbean Sea was land during the
Cretaceous and that at least some of this land was still land during
the Eocene.” F. M. Brown (private communication) states that current
geologic and oceanographic investigations, partly conducted by private
industry and not yet published, are yielding data which seem to indicate
that the Greater Antilles were indeed once connected with Central
America. At present, the entire question should be regarded without
prejudice as on open one.
182
FOX
J. Rri. Lcpid.
Schuchert ( 1935) thought that the Greater Antilles were connected
by dry land with Central America along what is now the Nicaraguan
swell — a high, wide hump of the sea floor running from Puerto
Rico, Hispanola and Jamaica to Honduras and Nicaragua — and by
a similar connection between Yucatan and western Cuba. He suggested
that the entire island group was part of the Central American mass
as recently as the Oligocene and th't duuring Miocene time Jamaica,
Hispanola and Puerto Rico were connected sporadically with the con-
tinent but that Cuba remained insular. Perhaps Schuchert was right.
Proponents of rafting have a second line of defense: the belief
that only the geography and climates of late Pliocene and of Pleistocene
times exerted significant influence on the distribution of modern
faunae. This belief is based on study of vertebrates, and nothing in
the present paper is intended as contradictory to the conclusions of
students of that phylum. The enviably full fossil record has permitted
mamalogists to reconstruct a reasonably complete picture of mammalian
evolution during the Tertiary. Oligocene time, for example, need not
be considered in connection with the distribution of most mammals,
since modern mammal species were not yet in existence.
Zoogeographers working with invertebrates, particularly with groups
like Lepidoptera for which there is almost no significant fossil record,
must rely upon principles of evolution and zoogeography developed
by mammalogists. "The fundamentals of invertebrate distribution are
substantially the same as those affecting vertebrates . . . The salient
difference is timing” (Miskimen, 1961). Jeannel (1949) pointed
out that, "the antiquity of insects becomes evident when compared
with that of vertebrate groups. The Tertiary and Quaternary lasted for
30 million years^ and comprised the entire evolution of the placental
mammals; but at the beginning of the Tertiary, most insect genera
already existed much as they do today.” According to Vandel (1949),
"The most evolved orders of insects (Lepidoptera, Diptera, Hymen-
optera) made their appearance during the Mesozoic epoch. At the
beginning of the Tertiary, the living entomological fauna [of the
world] was definitely constituted and thereafter underwent no important
alteration.” To evaluate insect distribution it is necessary to consider
events of the entire Tertiary, not just the effects of recent glaciations.
There is no doubt that the insect fauna of the north temperate
zone was profoundly affected by Pleistocene glaciation; the extent of
influence of glaciation on tropical and subtropical faunae is less clear
and in many regions may have been negligible. In all situations there
is a basic difference between the effect of the Pleistocene upon mammals
and its effect upon insects: mammals were evolving, insect' had already
evolved modern genera and species. Recent glaciation superimposed
certain geographic adjustments on pre-existing distributional patterns
of insects, but created the distributional patterns of modern mammals.
^Today we know the time estimate should be much longer.
2{}):I73-I84, 1963
ANTILLEAN ITHOMIIDAE
183
The Ithomiidae is one of the most primitive families of the Nym-
phaloidea (Fox, 1956) and must be of considerable antiquity. The
fact that one genus (Tellervo) is found in the East Indies while the
rest are neotropical suggests that the family must have been in existence
at the opening of the Tertiary. Even though Hymenitis is one of
the more specialized genera of the family, it probably was in existence
in Oligocene time. Thus if Schuchert’s connection between the Greater
Antilles and Central America during Oligocene or Miocene actually
existed, it would have provided a distributional route to the Greater
Antilles for these butterflies.
An objection to land route distribution — the same objection that
arises to rafting — is the question of why only certain Hymenitis
species are found on the islands. This is a question which cannot be
answered definitely, no matter which method of dispersal is postulated,
but it is easier to imagine that Hymenitis managed to survive while
other genera became extinct than it is to imagine that Hymenitis alone
of all the ithomines managed to beat the odds and find its way to the
islands from Central America.
SUMMARY
1. The only Ithomiidae found on the Antilles are Hymenitis
cubana on Cuba, H. d. diaphana on Jamaica and H. d. qtiisqueya
(herein described) on Hispanola.
2. The affinities of these Antillean ithomines are with certain
Central American Hymenitis. All are transparent-winged inhabitants
of the humid forests, shunning sunlight and open situations; all are
found from 1000 to 7000 feet above sea level.
3. Distribution by rafting is unlikely because of the direction of
sea and air currents and because of the habits and fragility of Hymen-
itis. If rafting ever distributed ithomines, the fauna of Venezuela and
the Guinas and the hardier sun-loving should be represented on the
islands.
4. Alternatively, these ithomines may have reached the Antilles
during early Tertiary time if, as Schuchert and others thought, the
islands once were continuous with Central America.
REEERENCES CITED
ANONYMOUS, 1807. Allg. Lit. Zeitg., Halle [Jena], 2. \fide Hemming, 1934]
AVINOFF, A. and N. SHOUMATOFF, 1946. An annotated list of the butter-
flies of Jamaica. Ann. Carnegie Mus., 30: 263-295.
BATES, M., 1935. The butterflies of Cuba. Bull. Mus. Comp. ZooL, 78: 65-258.
DARLINGTON, P. J., JR., 1957. Zoogeography: the geographical distribu-
tion of animals. John Wiley & Sons, Inc., New York.
DRURY, D., 1773. Illustrations of natural history, 2. London.
FOX, R. M., 1956. A monograph of the Ithomiidae. Part I. Bull. Amer.
Mus. Nat. Hist., Ill: 7-76.
184
FOX
J. Res. LePiJ.
GUNDLACH, J., 1881. Contribucion a la entomologia Cubana. Havana.
HEMMING, F., 1934. The generic names of the holarctic butterflies. British
Museum (Natural History), London.
HERRICH-SCHAEFFER, G. A., 1862. Schmetterlinge aus Cuba. Corresp.-
Blatt zoo. -min. Ver. Regensburg, 16: 118-120.
HtiBNER, J., 1816-1826. Verzeichniss bekannter Schmetterlinge. Augsburg.
JEANNEL, R., 1949. Les Insectes Fossiles. in P.-P. Grasse, Traite de Zoo-
logie, 9: 18-85. Masson et Cie, Paris.
MATTHEW, W. D., 1915. Climate and evolution. Reprint 1939, New York
Academy of Sciences.
MISKIMEN, G. W., 1961. Zoogeography of the coleopterous family Chauli-
ognathidae. Syst. Zool., 10: 140-153.
SCHUCHERT, C., 1935. Historical geology of the Antillean-Caribbean region.
John Wiley & Sons, Inc., New York.
SIMPSON, G. G., 1940. Mammals and land bridges. Jour. Washington Acad.
Sci., 30: 137-163.
1953. Evolution and geography. Oregon State System of Higher
Education, Eugene, Oregon.
WOODRING, W. P., 1954. Caribbean land and sea. Bull. Geol. Soc. Amer.,
65:720-730.
VANDEL, A., 1949. Generalities sur les Arthropodes. in P.-P. Grasse, Traite
de Zoologie, 6: 79-158.
Journal of Research on the Lepidoptera
2(3:) 185-200, 1963
ll-K) W. Orange Grove Are., Arcadia, California, U.S.A.
© Copyright 196'i
OVIPOSITIONAL PREFERENCE TESTS WITH PIERIS
WILLIAM HOVANITZ AND VINCENT C.S. CHANG
California Arboretum Foundation, Arcadia, California, and
Los Angeles State College, Los Angeles. California
Previous reports by us (Hovanitz and Chang, 1962, 1963)
have described the results of larval selections of various food plants.
The present paper continues this series of reports by describing the
selections made by the female adults on various food plants for the
purpose of egg laying. A similar type of study has been made by
Takata (1961) using Pieris rapae crucivora, though with somewhat
different results. This study will be mentioned below.
METHOD
The ovipositional tests were made in marquisette-covered cages,
18" X 18" X 24" tall. The plants used in each test were living, potted
plants grown in the glass house. In any given test, the plants selected
were all of about the same size and were relatively small so as to fit
in the dimensions of the cage size without severe crowding. The
differing shapes, growing habits and leaf sizes of the plants used
were variables which could not be controlled. There is no doubt that
these factors might have some influence on egg laying of the adult
females.
The females were placed in the cages with the respective plants
and allowed to mate and to oviposit. The eggs were counted periodically
for the totals indicated in the tables. The females used in each cage
varied in number from five to fifteen but the average was between
five and ten. These were fed honey and water solution in the mornings
during the course of each test.
RELATIONSHIP BETWEEN OVIPOSITIONAL PREFERENCE OF
THE FEMALE AND THE FOOD PLANT STRAIN OF ITS LARVA
A relationship is apparent between the ovipositional preference of
the adult female and the food plant eaten by its larva. This relationship
may be seen in table 1 in which strain A, Pieris napi, laid more eggs
on its ancestral food plant Dent aria than on any of the other plants
’Aided by a grant from the National Science Foundation, Washington, D.C.
185
Table 1. Ovipositional selections by adults of Pieris
186
HOVANITZ AND CHANG
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OVIPOSITIONAL TESTS
187
tested. Over forty percent were laid on Dentaria, about thirty-two
percent on black mustard ( Brassica nigra ) , about seventeen percent on
kale and less than one percent on Isomeris, out of a total of 259-
This relationship is further apparent in the data of strain B, Pieris
protodice ( table 1 ) . A strain of this species from Mono Lake, Calif,
where the native food plant is Lepidium, laid forty-one percent on
Lepidium, about twenty-four percent on mustard, about one percent
on kale and thirty-four percent on Isomeris, out of a total of 1295.
The high percentage on Isomeris is interesting in view of the fact that
Isomeris does not exist in Mono Lake area.
Another strain of Pieris protodice from Laguna Canyon, Orange
county, Calif., strain C, where the food plant is black mustard, gave
different results. Females of this strain laid nearly sixty-eight percent
of their eggs on black mustard, seven percent on kale, less than one
percent on nasturtium and twenty-five percent on Isomeris. Un-
fortunately, there was no Lepidium available for testing with these
females.
The two tests with Pieris protodice show a quantitative relationship
between the selection by the females and the odor of the plants selected.
Excluding naturtium, the plants may be arranged in the order Isomeris,
Lepidium, mustard and kale in the direction from strong odor to weak
odor. Females with Lepidium ancestry show a higher selection for
plants with a stronger odor, and plants with a mustard ancestry show
selections favoring a decreased odor requirement. In Table 1, strain
B (Lepidium ancestry) shows a selection toward Isomeris of 33 com-
pared with 25 for strain C (mustard ancestry). Likewise, strain B
shows only one percent toward kale as compared with seven percent
for strain C. The increase in selection of the mustard strain toward
mustard (68 percent as compared with 23 percent) is therefore not
the whole picture, as comparable changes occur also with the other
plants.
CHANGE OF OVIPOSITIONAL PREFERENCES BY FOOD
PLANT CHANGES OF THE LARVAE
Changing the food plants fed the larvae has the direct effect of
changing the plants selected by the adult females for egg laying. Table
2 shows the effect of splitting strain C ( table 1 ) into two strains,
one fed on kale (D) and the other fed on mustard (E). These were
grown for two generations on the respective food plants.
The results indicate an increased selection for kale by the kale
strain, 29 percent as compared with 7 percent. The strain kept on
mustard (E) did not vary greatly from the selections made by the
original parents strain (C).
Table 3„ Ovipositional selections by Pieris rapae. parent strains, and F^.,
Larvae always fed on food plant of female parent. Parent strains, 10 or more
generations on a particular plant.
188
HOVANITZ AND CHANG
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OVIPOSITIONAL TESTS
189
A similar relationship has been found with Pieris rapae (Table 3).
A strain kept for 10 or more generations on mustard in the laboratory
was compared with a strain grown an equal length of time on kale.
The mustard bred strain showed a selection of 87 percent on mustard
as compared with a selection by the kale strain of only 19 percent
on that plant; the kale strain adults selected kale 65 percent of the
time as compared with only 2 percent on kale from the mustard strain.
Pieris rapae, which have been switched over from kale or mustard
to Isomeris, also show an increased preference for the latter plant
but the selections take a lot longer period of time to become effective.
It has previously been shown that larvae of Pieris rapae grown on kale
for the first time have a high mortality rate approaching the lethal
condition. However, after a number of generations they are able to
survive satisfactorily on the new food plant. Ovipositional tests (Table
4) of Pieris rapae reared on Isomeris show that adults from the second
generation of larvae fed on Isomeris show little change in their selec-
tion. Most eggs are still laid on mustard with a high proportion on
kale and nasturtium. In the fourth generation however, there are 24
percent of the eggs laid on Isomeris and a corresponding drop in those
laid on nasturtium. A more complete series of tests of this sort is
shown in Table 5 which will be discussed below.
GENETIC CHANGE OF OVIPOSITIONAL PREFERENCES AS
SHOWN BY INTERSTRAIN CROSSES
The genetic significance of the changes in ovipositional preferences
as created by larval feeding habits are shown by crosses involving parent
strains, one of which has been adapted to kale and the other to mustard
(Table 3). The parent strains have been grown for ten or more gen-
erations on their respective food plants and show a high ovipositional
selection for their own larval food plants ( fig. 1 ) .
The F] of the cross kale-strain female X mustard-strain male was
grown on kale and the reciprocal cross was grown on mustard. Ovi-
positional preferences in the first of these two crosses indicate that
the El adults, even though fed on kale as larvae, did not select kale
to the extent of the kale parental strain. There was a drop in selection
of kale from 65 percent to 30 percent. The increase appears to have
all gone to the selection of nasturtium ( fig. 2 ) .
In the reciprocal cross, the Fi larvae were grown on mustard. The
adults from these larvae dropped in their selection of mustard from
87 perecent to 35 percent. The increase in this case was not only
toward nasturtium but also toward kale.
It is clear from these crosses that the genes for the inheritance of
food plant preferences by the adult for ovipositional purposes is trans-
mitted through the male as well as through the female. Indications
PERCENT SELECTION PERCENT SELECTION
190
HOVANITZ AND CHANG
]. Res. Lepid.
Fig. 1. Oviposition selection by females of Pieris rapae for the plant on
which the strain has been maintained for more than 10 generations.
PERCENT SELECTION PERCENT SELECTION
OVIPOSITIONAL TESTS
191
2(3);IS5-200, 196)
Fig. 2. Oviposition selection by females of Pieris rapae produced by
crossing mustard and kale strains to obtain the Fi.
Table 5. Ovipositional selections by adults of P. rapae grown on nasturtium
given number of generations, (former plant, kale)
192
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Table 7^ Ovipositional selections by adults of P, rapae grown on kale for
given number of generations^ (former plant, kale)
2(}):lS5-200, 196}
OVIPOSITIONAL TESTS
193
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194
HOVANITZ AND CHANG
J. Res. LepiJ.
that these traits may be maternally inherited are therefore not wholly
if at all true.
The of the mustard female Fi and the reciprocal shows a some-
what similiar selection pattern as the F^ with the exception that the
selection of nasturtium has been reduced in favor of both mustard and
kale ( fig. 3 ) .
RATE OF CHANGE OF OVIPOSITIONAL PREFERENCES OVER
A PERIOD OF SEVERAL GENERATIONS
The rate of change of ovipositional preferences during the course
of several generations has been studied in three series, one in which
the larvae have been bred on nasturtium, one on Isomeris and one on
kale. These data are indicated in tables 5, 6 and 7.
Nasturtium is not a normal food plant of Pieris rapae, but colonies
of the insect have become adapted to the plant both in Europe and
in the United States. Larvae from a strain previously fed on kale were
transferred to nasturtium and maintained on that plant for eight gen-
erations. Adults from some of these generations were tested for their
preferences toward kale, mustard, nasturtium and Isomeris. The nor-
mal selection of kale strain adults to Isomeris is about 10 percent
( Table 3 ) • The first generation after the larvae were fed nasturtium,
the selection went up to 20 percent, the third generation 38 percent,
the fifth generation 45 percent, the seventh generation 34 percent and
the eighth generation 37 percent. The lack of any change above 34
percent may mean that this is the best possible selection that can be
obtained after any amount of selection.
It is significant that during the course of these selections for
nasturtium, selection for kale went progressively down and that at
the same time selection for Isomeris went progressively up. As has
been previously pointed out, this result may be due to the stronger
odoriferous substances present in nasturtium and Isomeris, absent in kale.
Isomeris is not known to be a wild food plant for Pieris rapae
anywhere. Larvae are, however, able to survive on this plant, especially
after conditioning. Larvae from a strain fed previously on kale were
trensferred to Isomeris and maintained on this plant for eleven gen-
erations. The normal selection of kale strain adults to Isomeris is
about 2 percent (Table 3). The first generation of adults from larvae
fed on Isomeris selected this plant 4 percent of the time, the third
generation 12 percent of the time, the fifth generation 8 percent of
the time, the seventh generation 32 percent of the time, the ninth
generation 27 percent of the time and the eleventh generation 19
percent of the time ( fig 5 ) . A significant increase is apparent, though
the variation is somewhat great, and this increase is additive for several
generations. It is of interest that the selection of Isomeris, though
PERCENT SELECTION PERCENT SELECTION
2n):ns.200, /96!
OVIPOSITIONAL TESTS
195
Fig, 3- Oviposition selection by females of Pier is rapae produced by
crossing the Fi individuals indicated in fig. 2.
196
HOVANITZ AND CHANG
/. Res. Lcl)ii!.
increased, is never greater than that for kale and only comes close to
duplicating the selections for mustard.
Strains grown on kale for twelve generations, after originating from
wild individuals growing on mustard, were tested in the third ex-
periment. A mustard strain selected kale about two percent of the
time. After one generation on kale, the selection of kale was 48 per-
cent, after three generations 69 percent, five generations 69 percent,
seven generations 62 percent, eight generations 38 percent, nine gen-
erations 43 percent, ten generations 45 percent and twelve generations
60 percent (fig. 6). Again there is no doubt of the increased rate of
selection of the larval food plant by the adults for the first few gen-
erations though there are somewhat erratic results thereafter due to
unknown causes. In fact after three generations the selections of kale
were as good as is expected in a strain kept permanently on kale
(Table 3).
This experiment indicates that adaptation to kale is much faster
than to plants such as Isomeris and nasturtium, and is also much more
effective.
ADDENDUM
The following data were made available after the previous portion
of this paper was written. Adults of Pieris protodice were collected in
the vicinity of Palm Springs, in an area typical of the Coachella desert.
As a result it cannot be anticipated that the cruciferous plants typical
of coastal California were present. Instead, the bushes of Isomeris are
the most abundant plants of this group. These females were allowed
to oviposit in tests as the preceding, with the result that Isomeris was
the plant most commonly selected (59 percent) with mustard in
abundance (41 percent) (Table 8). These data compare with P,
protodice selections in an area where mustard is the wild food plant
as follows: Isomeris 25 percent instead of 59 percent, and mustard
67 percent instead of 41 percent.
DISCUSSION AND CONCLUSIONS
These results of ovipositional experiments illustrate the same events
as have been shown previously for larval selections of food plants,
namely that previous exposure to a plant is correlated with, the selec-
tion of that plant by the particular insect. This increase of selection
occurs not only in the first generation of exposure, but also after sub-
sequent generations of exposure. It is easier to change the selection of
Pieris toward kale than it is toward Isomeris or nasturtium. This is
true both with respect to the speed with which the change is made
as well as in the extent to which the change occurs.
2(}):I85-200, 196}
OVIPOSITIONAL TESTS
197
Fig. 4. The change in ovipositional selection by adults of Pieris rapae
after 8 generations of growing on nasturtium.
198 HOVANITZ AND CHANG /• LcpuI.
Fig. 5. The change in ovipositional selection by adults of Pieris rapae
after eleven generations of growing on Isomeris.
2(}):185-200, i96.i
OVIPOSITIONAL
TESTS
199
60-
%4 0 -
2 0
60
% 40 -
20 -
N I
Fig. 6. The change in ovipositional selection by adults of Pieris rapae
after twelve generations of growing on kale.
200
HOVANITZ AND CHANG
/. Res. LcpiiL
The heredity of the food plant selections has been tested by both
Fi and F2 crosses. The results indicate that the Fi selections are inter-
mediate between the parental selections and that the F2 is also, but
the latter tends to match the food plant on which the F2 larvae are
fed. An unknown cause accounts for a heavy increase in selection of
nasturtium in the Fi which is not so apparent in the F2. The gene
or genes for the food plant specificities is clearly carried both by the
males as well as by the females and there is no sign of maternal
inheritance, except a psuedo-maternal effect which is imposed upon
the progeny by the food directly eaten by them.
These results are directly opposed to those of Takata (1961) in
which he says: "If the larvae were reared with cabbage for successively
many generations, the emerged butterflies tended more and more to
avoid cabbage as their ovipositing plant, as the successive life on
cabbage was lengthened . . . The direction of the change was quite
inverse to that of the larval food preference, which was shown by that
the larvae preferred the host plant on which they and their ancestors
had been fed.” No reason can be given for the difference between
his results and ours, nor is their any clue as to what it might be with
the one exception: In our own experiments, more than two plants
were available both for larval selections as well as for adult selections.
It was possible in our experiments therefore for the adults to select
some plant other than the two being tested in case neither of the
original two were satisfactory. As an example of this phenomenon, in
the Fi of the mustard-kale strain crosses, the greatest proportion of
the adults selected nasturtium rather than either mustard or kale. Had
nasturtium not been available, the adults would have had to choose
between two plants neither of which was preferred. Something of this
sort might explain the results of Takata.
LITERATURE CITED
HOVANITZ, W. and V.C.S. CHANG. 1962. Three factors affecting larval
choice of food plant. J. Res. Lepid. 1(1) : 51-61.
1962. The effect of various food plants on survival and growth
rate of Pieris. /. Res. Lepid. 1(1): 21-42.
1963. Change of food plant preference by larvae of Pieris rapae
controlled by strain selection, and the inheritance of this trait. J. Res.
Lepid. 1(2): 163-168.
1963. The effect of hybridization of host-plant strains on growth
rate and mortality of Pieris rapae. /. Res. Lepid. 1 (2) : 157-162.
TAKATA, N. 1961. Studies on the host preference of common cabbage
butterfly, Pieris rapae crucivora (Boisduval). XII. Successive rearing
of the cabbage butterfly larva with certain host plants and its effect on
the ovipositional preference of the adult. ]ap. J, Ecology 11: 147-154.
Journal of Research on the Lepidoptera
2(3)201=203, 1963
© Copyright 196)
1140 W. Orange Grove Are., Arcadia, California, U.S.A.
NOTES ON THE EARLY STAGES OF
DREPANULATRIX MONICARIA (GUENEE)
(GEOMETRIDAE)
JOHN ADAMS COMSTOCK
Del Alar, California
The life history of this moth has been treated at some length
by Dr. Frederick H. Rindge in his classic "Revision of the Genus” in
1949. No illustrations were included other than of the genitalia.
I have recently reared this species, and made drawings of the egg,
first instar larva, final instar larva and pupa. The brief notes accom-
panying- these drawings will serve to supplement, and in a few par-
ticulars to amplify Dr. Rindge’s text.
A gravid female of Drepanulatrix monicaria was captured on May
22, 1962, in Del Mar, California. Numerous eggs were laid May 23 and
24. These were laid singly on their sides. They hatched June 3 and 4.
EGG: The egg is ovoid, the base flattened and the top regularly
rounded. It is 0.75 mm. high by 0.50 mm. wide on the average.
The accompanying drawing, (figure lA) shows it in an upright
posture on which I base the measurement of height and width, but
Dr. Ridge gives its measurement in relation to the normal prone posi-
tion, hence the disparity in our terms.
I could detect no "flattening” of viable eggs, nor any that were
"wedge shaped.”
When first laid, the color was a glistening greenish white. Subse-
quently it changed to mottled and streaked reddish orange of varying
intensities. The surface is covered with longitudinal ridges, approximately
24 in number. Each ridge carries a line of minute pearl-like nodules along
its edge. Between the ridges there are transverse lines or low ridges.
The base is covered by irregular hexagonal cells with raised walls. The
longitudinal ridges seem to fuse into a slightly pitted surface over the
rounded top, but no regular micropyle could be detected.
The egg is very similar to that of Drepanulatrix huhti, but is
smaller.
The young larvae exit from one end, the remainder of the shell being
left intact.
The recorded food plants for this species are Ceanothus thyrsiflorm
Eschs., C. macrocarpus Nutt., C. spinosus Nutt., and C. sorediatus Hooker
& Arnott. Our young larvae readily accepted the first named plant.
FIRST INSTAR LARVA: Length, 3 mm.
Head width, 0.30 to 0.33 mm. Color, glistening yellow, with a
201
202
COMSTOCK
/. Res. LepiJ'.
narrow dark margin on the edge of the posterior juncture with the first
body segment. Ocelli, black. There is no spotting on any portion. Setae,
short and colorless to white.
Body; ground color, light yellow-green, with a wide middorsal band
which is olive-green on the first two segments, shading into red-brown
caudally.
The first segment is narrower than the head and wider than the
second segment. The light green on this segment is slightly tinged with
yellow. The black papillae on the body are relatively smaller than those
on Drepanulatrix hulsti, but occupy much the same positions. They are
narrowly encircled at their bases by white, in strong contract to the wide
white circlets on D. hulsti.
The setae are black. A substigmatal band is usually present.
The first larval instar is shown in dorsal aspect on figure IB.
FINAL INSTAR: This is given in detail by Dr. Rindge in his
Revision. The wide variation in color and markings is particularly
noteworthy. No two examples are exactly alike. The color ranges from
a smoky gray (almost black and heavily spotted) to a rich spotted
green. Two dark examples have a discontinuous middorsal line of burnt
orange. The green and tan examples lack this, or show only slight
suggestions of it. All types are sprinkled with either light or dark spots.
The specimen I selected was probably in the early phase of the final
instar and not fully grown as it measured only 13 mm. It is pictured
in lateral aspect on figure 1C. In this example the head was yellow and
the body green, with white spots.
PUPA: Length, 10 mm. Greatest width, 3 mm. Head well rounded.
Eyes not prominent. Antennae and maxillae extending to wing mar-
gins. Abdomen tapering to cremaster, — the latter ending in a large
papillus which bears four recurved booklets each side of the median
line. Two of these, centrally placed, are relatively long. All arch laterally.
The color of the chrysalis is deep brown, with a slight tinge of
green on the wing cases in newly formed pupae. The surface texture
is glistening.
The pupa is shown in figure ID. An enlarged figure of the cremaster
is included on figure IE.
The first imago emerged July 23, 1962.
This moth ranges along the Pacific coast from southern Alaska to
the Mexican border, and perhaps into northern Baja California.
Dr. Rindge recorded three parasites, i.e. Chaetophlepsis sp., Patelloa
sp., and Apanteles sp.
REFERENCES
RINDGE, FREDERICK H. 1949. A Rev. of the Geometr. Moths formerly
assigned to Drepanulatrix. Lepidoptera. Bull. Am. Mus. Nat. Hist.
94: (59) pp. 233-298. Egg, larva, pupa, food pi.
McFarland, NOEL. 1959. in litt. l/21/’59. Food pi.
2f3:)20-20}, 1963
DREPANULATRIX MONICARIA
203
FIGURE 1. Early stages of Drepenulatrix monicaria (Guenee) A. Egg,
enlarged X 90. B. First instar larva, enlarged X 40. C. Mature larva on
twig of Ceanothus, enlarged X 10. D. Pupa, ventral aspect, enlarged X 10.
E. Cremaster, enlarged X 20.
Reproduced from water color drawing by the author.
204
COMSTOCK
/. Res. Lepid'.
BIOGRAPHICAL SKETCHES
COMSTOCK, JOHN ADAMS
[1373 Crest Road, Del Mar, California]
Born: Evanston, Illinois, Jan, 30, 1883. Son of John Adams Comstock, descendent of
William Comstock, from England, 1637, and Nellie Hurd, daughter of Judge Harvey B. Hurd,
one of the founders of Northwestern University.
M. A, (Hon, ): Occidental College, 1924
D. O, : College of Osteopathic Phys. and Surg. , 1945
D. Sc. (Hon.): College of Osteopathic Phys. and Surg., 1945
M, D. : California College of Medicine, 1962
D. Sc. ; California College of Medicine, 1962
Fellow: Roy. Ent. Soc. London; Sou. Calf. Acad. Sci. ; Inst. Amer. Genealogy; San Diego Soc,
Nat. Hist.
Selected List of Past Positions: Secretary, Chicago Entom. Soc., 1898-1903
Recorder, Entom, Soc., Chicago Acad. Sci., 1903-04
Taught Anatomy and Embryology, Coll, Phys. Surg., 1915-1920
Assist, and Dir. Southwest Museum, 1920-1926
Editor, Publ. Sou. Calif. Acad. Sci., 19 2 1-62
Act. and Assist. Dir. L, A. Co. Museum, 1928-
Director Science, L. A. Co. Museum, 1939-48.
Retired 1948
President Fellows, San Diego Soc. Nat, Hist.
1958; Lepid. Soc., 1962
Member, Board of Directors, Nat. Hist. Mus.
San Diego, 1957
Interests: Butterflies of California (book, 1927)
Geneology (book "History and Geneology of the
Comstock family in America, " 1949)
Life Histories of Lepidoptera
Publications: 194+ papers mostly illustrated
life histories, 1902-1963
Journal of Research on the Lepidoptera
2(3) :205-223, 1963
1140 W . Orange Grove Avc., Arcadia, California, U.S.A.
© Copyright 196}
THE ORIGIN OF A SYMPATRIC SPECIES IN COLIAS
THROUGH THE AID OF NATURAL HYBRIDIZATION
WILLIAM HOVANITZ ^
[continued from volume 1, page 274]
IV POPULATION ANALYSES FROM SIGNIFICANT REGIONS
Analysis of the variation of Colias hecla. Colias nastes and Colias
hoothi (where present) has been made at a large number of stations
throughout arctic North America and at one station in northern
Sweden. The North American stations are shown on the map (fig, 10)
drawn for this purpose. Many of the stations shown are taken from
the map drawn for the "Northern Insect Survey” of the Canadian
Government and made available for this use by Dr. T. N. Freeman.
In addition to the localities which are taken from that map, some
stations are shown which represent samples made by others. There
are included a sample from the Meade River, Alaska made available
by Dr. John Garth, a sample from Coppermine, Northwest Territories,
collected by the author in 1947, a sample from Danske Island in the
Hudson Bay made available by the Danish National Museum, Copen-
hagen, and a sample from Coral Harbour, Southampton Island, collected
by Dr. G. M. Sutton and made available by the Garneigie Museum,
Pittsburgh.
The analyses of the populations has been made by the two char-
acters indicated in section III in this series, namely, by applying a
numerical value to the grade of border pattern displayed by each
specimen, and also by applying a numerical value to the grade or
orange pigmentation displayed by each specimen. These two values
for each specimen are then plotted on a scatter diagram, which gives
a fair idea, on perusal, of the condition of the variation of the three
species in any one locality. As the females all show the same border
pattern for both species, this sex was rated by color only. These dia-
grams are indicated as figures 11 through 42. The method of grading
the insects is described in section III and is illustrated in color in
figures 8 and 9.
205
Fig. 9. Series of Colias, mostly identifiable as C. boothi, showing by means
of a graded series, nine steps in the orange color development on the fore
wings of the male from the yellow at the top left to the full orange at the
second to the bottom right. Grades are designated as 0 through 4 on the
left column and 5 through 9 on the right column. The specimen on the
bottom right does not enter the series but is shown to illustrate a variation
in which the fore wings are orange and the hind wings yellow. All specimens
shown are males. All are from Coppermine, Northwest Territories, Canada,
July 12-19, 1947, W. Hovanitz coll, except (1) grade 0, top on left column.
Repulse Bay, Northwest Territories, Canada, Aug. 2, 1950. P. F. Bruggemann,
(2) Grade 8, fourth from top on right column. Repulse Bay, N, W. T.,
Canada, July 27, 1950. J. E. H. Martin.
EUREKA
2('3jr205-223, 1963
SYMPATRIC SPECIES
207
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tions of Colias which have been graded for two characters.
COPPERMINE ... DANSKE
2(3):20^-22J, 1963
SYMPATRIC SPECIES
209
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SYMPATRIC SPECIES
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215
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219
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224
HOVANIT2
/. Res. LepiJ.
BIOGRAPHICAL SKETCHES
TILDEN, JAMES WILSON
[125 Cedar Lane, San Jose, Calif^ ]
Born: Philo, Calif, , 1904
Married: 1943, 3 children
A, B, : San Jose State College, 1942
M, A,, PhD,: Stanford University, 1948
Associate Editor: Lepid, News, 1959
Professor of Entomology: San Jose State College, 1948-,
Consultant: Santa Clara County Health Department. 1955-59
Contributor to many scientific journa]
Interests: Ecology, Distribution and
Migration of Lepidoptera, espe-
cially Hesperiidae and Lycaenidae,
Teaching: Advanced General Entomo-
logy, Biological Control, Forest
Entomology,
Journal of Research on the Lepidoptera
2(3) :225-228, 1963
1140 W. Orange Grove Arc., Areaclia, California, U.S.A.
© Copyright 1961
A RUBBER STAMP METHOD EOR PRODUCING
SPECIMEN LABELS
PHILLIP A. ADAMS
Orange^State College, Fullerton, California
The method described below has proved a speedy and con-
venient means of producing insect labels, and appears equally adapt-
able for labels of larger size. Anyone capable of lettering neatly with
a crow-quill pen should have little difficulty in mastering the tech-
nique. Some practice probably will be required before best results
consistently are obtained. These directions are the result of much
experimentation, and should be followed meticulously. Brand names
of several materials are given because other brands have not proved
so satisfactory; substitutions should be avoided.
In view of modern advances in techniques of reproduction, prep-
aration of the mold by hand engraving may appear retrogressive. How-
ever, this method offers several advantages. No expensive equipment
is required, and all the materials are easily obtainable. There is no
necessity for setting up or distributing type, nor is there a press to
clean. The equipment may be taken into the field. Most important,
the method is extremely rapid. Maximum efficiency is attained for
series of eight to fifty labels.
Ross scratchboard makes an ideal mold for insect labels. The thin-
ness of the plaster surface limits the stamp to low relief; consequently
letters must be made small and close together. For larger lettering, as
is used in bird and mamm.al labels, where deeper engraving is required,
thin tablets of plaster of paris are satisfactory. Wax is not suitable as
a mold, since the shavings clog the impressions of the letters.
Lettering is cut with a dissecting needle, shortened to 8 mm and
resharpened. The needle should be held vertically, and gentle, even
pressure used. Letters with closed loops must be executed with par-
ticular care to prevent the centers of the loops from breaking out.
Because of the low relief in stamps with fine lettering, blank spaces
tend to print as smudges. If spaces cannot be avoided, lines should be
ruled through them ( fig. 1 ) . Engraving these labels is so rapid that
a separate mold easily may be made for each date, thus eliminating
the necessity of annotating individual labels with pen.
Letters need not be cut deeply to print well. Avoid cutting
through the plaster coating of the scratchboard to the paper beneath;
225
226
ADAMS
J. Rea. Lrpiif.
if this occurs, the rubber often pulls up flakes of plaster as it is sep-
arated from the mold. This sticking may be prevented by treating the
scratchboard with silicone mold-release, or with polypropylene glycol
(hydraulic brake fluid). Ordinarily such treatment is not necessary,
unless the board* has been coated with ink, but it greatly improves the
working qualities of the scratchboard.
Make a frame by cutting in a piece of thin, waxed cardboard, a
rectangular opening large enough to allow a space of at least 4 mm
surrounding the engraved label, when the frame is placed over the label.
This frame will depress the face of the stamp surrounding the label,
so that only the portion included in the opening will print. Using the
frame as a guide, incise a border no farther than 1 mm from the letter-
ing, and slightly deeper ( fig. 1 ) . The space between this border and
the cardboard frame is then filled with parallel engraved lines ( fig. 2 ) .
This border area supports the face of the stamp, preventing smudging,
and allowing more pressure to be used in printing, without wearing
down the letters so rapidly. The wider it is, the better will be the
quality of the labels. The printed border is trimmed away from the
finished labels.
A stamp handle is made by cementing a rectangle of glass or metal,
10 X 30 mm, to the end of a large cork (fig. 5). Over this is placed
a 20 X 70 mm strip of wire gauze, the ends of which are bent back
and taped to the sides of the cork.
To make the stamp, a small pellet of Eberhard Faber kneaded
rubber is laid over the mold ( fig. 3 ) . The cardboard frame is steadied
with the left hand while the stamp handle is pressed down firmly into
the rubber, and removed with a slight rocking motion. This should
result in a thin layer of rubber on the face of the stamp handle, ex-
tending about 3 mm beyond the impression of the frame, and bearing
a perfect impression of the mold ( fig. 4 ) . If too much rubber has
been used, the edges should be pressed down to prevent their printing.
The stamp pad is a piece of one fourth inch foam neoprene, such
as is used in skin diver’s wet suits, covered with a tough piece of
plastic film, taped in place. After each use, the plastic is stripped off
and discarded. Rigid surfaces, such as with glass, are not suitable for
ink plates, since they do not ink the stamp evenly, and their lack of
resilience causes the stamp to wear down rapidly. Cloth stamp pads
are too coarse for insect labels, and cannot be used with oil ink.
The ink is spread with a brayer made from a piece of coat-hanger
wire, with a 40 mm length of tygon plastic or smooth rubber tubing
for a roller. The tubing may be discarded after each use. If shaped
as shown in figure 6, the brayer may be set down without the inked
roller touching the surface.
Either ordinary printers ink or oil linoleum block-printing ink
may be used. It should be spread out quite thin, and frequently rolled.
If allowed to become tacky, the ink pulls rubber from the face of the
stamp.
2(}):225-228, 196}
SPECIMEN LABELS
227
Fig. 1. Incising the inner border around the label, using a cardboard
frame as a guide. Fig. 2. The completed label mold, engraved in scratchboard.
Note wide engraved border. Fig. 3. Cardboard frame and kneaded rubber
pellet in position over mold. Fig. 4. The stamp, formed for forcing the rubber
into the mold with the stamp handle. Edges of rubber being pressed down
to prevent smudging. Fig. 5. The stamp handle. Wire gauze is held in place
with a few turns of tape. Fig. 6. Brayer. Fig. 7. Printing pad with guide
for aligning labels.
When inking and printing, hold the cork gently between the thumb
and third finger, with the index finger on the top of the cork. Touch
the stamp to the inked surface, release the thumb and third finger,
and press lightly with the index finger. Print with the same motion.
Very little pressure is required; delicate handling produces clean im-
pressions and prevents premature deterioration of the stamp. Two
labels usually can be printed each time the stamp is inked.
Any medium weight paper may be used, so long as the surface
is smooth. For very small lettering, a coated paper, such as is used
to line insect boxes, is ideal. A sheet of foam neoprene under the paper
results in better impressions and prolongs the life of the stamp. Straight
rows of labels may be produced using a margin guide as shown in
228
ADAMS
/. Rea. Lepid.
figure 7. With this device printing is slightly more difficult, but
cutting out the labels is greatly facilitated.
After about fifty impressions, a kneaded rubber stamp usually will
require renewal. Remove the ink with scrap paper, peel off the rubber,
reshape it, and take another impression from the mold. Discard the
rubber when it absorbs so much ink as to lose its springiness, or be-
comes sticky.
Permanent stamps can be made with a variety of materials. Silicone
rubber has been used with good success. The mold should be lubri-
cated with green soap or silicone before engraving. A long curing
time — twenty-four hours or more — may be required. If difficulty
is experienced peeling the silicone from the mold, the scratchboard
can be dissolved in hot potassium hydroxide solution. Kneaded rubber
is much easier to use. Results with permanent stamps have not been
sufficiently better to make the extra trouble and delay seem worth
while.
Mr. Richard Fall, of Bio Metal Associates, has given encourage-
ment and valuable suggestions, and has kindly assisted in procuring
materials used in developing the technique. I am gratefuly to R. B.
Cowles for his comments on the manuscript.
Journal of Research on the Lepidoptera
2(3) :229-239, 1963
1140 W. Orange Groic Ate., Arcadia, California, U.S.A.
© Copyright 196)
GENERIC OR SUBGENERIC NAMES CLOSELY RELATED
TO ARGYNNIS
PADDY B. McHENRY
1032 E. Santa Anita Are.. Burbank. California
A REVIEW WAS MADE of the generic and subgeneric names that
have been applied over the world to the Argynnid butterflies as an
incidental to preparing a list of the generic and specific names that
have been used for the Neartic Argynnids. This paper is a summary
of that study.
The names are listed in alphabetical order. Synonyms (where the
types are the same) are listed alphabetically but under the names
having priority. As no study of the biological relationships of this
group of insects has been made, no attempt has been made to indicate
preferential use of the names as genera, subgenera, or if the names
should be used at all.
Various authors have dealt at length with the generic arrangements
for this group of butterflies, for example Reuss (1926, [1927]) : War-
ren (1944, 1955); Dos Passos and Grey (1945); and Warren, Dos
Passos and Grey (1946). These authors should be consulted for bio-
logical diagnosis of certain species and for definitions and subjective
analyses of the genera.
Dos Passos and Grey (1945) associated the genus Euptioeta
Doubleday (1848) with the Argynnid butterflies but Warren, Dos
Passos and Grey (1946) dropped this generic name from that consid-
eration.
A chronological history of the names of generic (or subgeneric)
rank proposed for the Argynnid butterflies is summarized as follows:
1758. Linnaeus (1758) applied the name Papilio to all butter-
flies. Argynnids were given this name from 1758 through the first part
of the 1800’s.
1777. Scopoli erected the name Argyreus for a group of assorted
butterflies which included some Argynnids.
[1806]. Hiibner erected the name Dry as for paphia Linnaeus in an
"unpublished” work.
1807. Fabricius erected the name Argynnis which has been the
most generally accepted name for the species of this group since its
origin.
229
230
McHENRY
J. Res. LepiJ.
[1819]. Hiibner recognized Argynnis Fabricius and erected four
additional generic names for some of the Argynnids; Acidalia, Ai.rgyron-
ome, Brenthis and Issoria. Recognition of these names was slow and
much confusion was caused since no type species were originally des-
ignated. The name Brenthis appears to have been generally misused.
1820. Billberg essentially applied the name Argyrea to the Argynnid
butterflies. Little or no use has been made of this name.
1851. Nordman erected the name Damora in connection with de-
scribing a new species. Very little use has been made of this name also.
1872 and 1875. Scudder erected the names Speyeria and Semno-
pysche for certain Nearctic species. These names have only recently
been resurrected by Dos Passos and Grey for generic {Speyeria) and
subgeneric {Semnopysche) use. Editorial policy in one journal has
aided in the popularization of the name Speyeria as a generic name.
1900. Moore in his "Lepidoptera Indica” erected the names Boloria
and Rathora. Reuss helped to put the name Boloria to common use.
1920 to 1928. Reuss erected the following names for Argynnid
genera or subgenera: Eudaphne, Eudryas, Clossiatia, Fahriciana, Kilken-
thaliella, Mimargyra, Mesoacidalia, Mesodryas, Neoacidalia, Proacidalia,
Proclossiana, Prodryas, Profabriciana, Prokukenthaliella, Protodryas,
Pseudorathora, and Yramea. These names included some homonyms
and synonyms.
1942. Warren erected the name Pafidoriana in connection with his
studies.
1943. Hemming proposed Childrena to replace Endryas Reuss which
w^as preoccupied.
1944. Bryk erected the name Chilargynnis for cytheris Drury ap-
parently being unaware of Reuss’ Yramea with the same type.
The use of these names varies greatly depending upon ( 1 ) the
preference by specialists for large or small genera, or ( 2 ) whether the
group as a whole is considered a subfamily or a genus. At the one
extreme, each species of the group might be considered under a different
generic name and the group then is called Argynninae; at the other ex-
treme, all species of the group might be considered under one generic
name and the group is then called Argynnis.
2(}):229.2i9, 1963
ARGYNNID NAMES
231
LIST OF NAMES USED FOR GENERIC
OR SUBGENERIC PURPOSES
ARGYNNIS FABRICIUS.
Type: paphia Fabricius = paphia Linnaeus,
ARGYREA BILL BERG. ”
Type: paphia Billberg = paphia Linnaeus,
DRYAS HUBNER (in Tentamen).
Type: paphia Hubner = paphia Linnaeus.
M E S O D R YAS REUSS. “
Type: paphia Linnaeus,
ARGYREUS SCOPOLI.
Type: niphe L. = hyperbius Johansson [Linnaeus ?].
A CIDALIA HUBNER.
Type: niphe L. = hyperbius Johansson [Linnaeus?].
MIMARG y“10~REUSSI “
Type: hyperbius Johansson [Linnaeus ?], Later named niphe by
Linnaeus.
ARGYRONOME HUBNER.
Type: laodice Pallas.
E U D A P H NE REUSS.
Type: laodice Pallas.
B GLORIA MOORE.
Type: pales Schiffermuller [Denis and Schiffermuller],
BRENTHIS HUBNER.
Type: hecate Wien. Verz, [Denis and Schiffermuller],
CHILDRENA HEMMING.
Type: children! G. Gray.
E U D R Y A S REUSS.
Type: children! G. Gray.
CLOSSIANA REUSS.
Type: selene [Denis and Schiffermuller],
DA MORA NORDMANN.
Type: paulina Nordmann = sagana Dbdy, and Hew,
FABRICIANA REUSS.
Type: niobe Linnaeus,
ISSORIA HUBNER.
Type: lathonia Linnaeus.
R A T H O R A MOORE.
Type: lathonia Linnaeus.
KUKENTHALI ELLA REUSS,
Type: gemmata Butler.
MESOA CIDALIA REUSS.
Type: aglaia L. = charlotta Haworth.
NEOA CIDALIA REUSS. ““ ”
Type; cybele Cramer [Fabricius],
PANDORIANA WARREN.
Type: maja Cramer = pandora Schiffermuller [Denis and Schif-
fermuller].
PROACIDALIA REUSS.
Type: clara Blanchard.
PROCLOSSIA NA REUSS.
Type: aphirape Hubner = eunomia Esper.
232
McHENRY
/. Res. LepiJ.
PROFABRICIANA REUSS,
Type: jainadeva Moore.
prokukenth'aliella REUSS.
Type: excelsior Butler.
PROTODRYAS REUSS. '
Type: kamala Moore.
P R O D R Y A“s“¥eUSS.
Type: kamala Moore.
PSEUDORAT HORA REUSS.
Type: isaeea Doubleday (Nomen nudum) = issaea Moore.
SEMNOPYSCHE SCUDDER. ”” ' “
Type: diana Cramer.
SPEYERIA SCUDDER.
Type: idalia Drury.
YRAMEA REUSS^
Type: cytheris Drury.
CHILARGYNNIS BRYK.
Type: cytheris Drury.
DATA FOR THE NAMES
ARGYNNIS FABRICIUS. 1807 [before 19Dec.]\ In K. Illiger. Magazin
fur Insektenkunde 6: Z83, no. 19. Includes among others "Pap.
Paphia" which is considered to be of Linnaeus.
Type. P[apilio]. N[ ymphalis], [Phaleratus] paphia Linnaeus. 1758.
Syst. Nat. 10th. Ed. 1: 481, no, 138.
Type Selection. Latreille. 1810. Consid, Gen. Anim. Crust. Arach.
Ins., p. 440. Says in a s ection devoted to types of genera, after
having established that Argynne = Argynnis: "Argynne. Argyn-
nis paphia. Fab. ; ejusd. , melitaea cinxia". As used by Lat-
reille paphia Fab. is considered to be paphia Linnaeus,
Note. Hemming. 1958. Official List of Generic Names in Zoology.
1st. Install.: 69, no. 609. Argynnis is included on Official List
and has preference over Argyreus Scopoli.
ARGYREA BILLBERG. 1820. Enumeratio Insectorum in Mus eo Gust.
Joh. Billberg; p. 77, Includes among others "paphia" which
is considered to be of Linnaeus,
Type, P[apilio], N[ ymphalis], [Phaleratus] paphia Linnaeus. 1758.
Syst, Nat. 10th. Ed. 1: 481, no. 138.
Type Selection, Hemming. Sept, 1933, Entomologist 66 (844): 197.
Says of Argyrea: "Type = Argyrea paphia Linn, "
DRY AS HUBNER, [1806]^. Tentamen Deter. Digest, Demon, Singul.
Stirp. Lepidopterorxim; p, [1], no. IV of Tribus 1 listed un-
der Phalanx 1.
Type, P[ apilio], N[ ymphalis], [ Phaleratus] paphia Linnaeus, 1758.
Syst. Nat, 10th. Ed. 1: 481, no. 138.
Type Selection, Hubner, As above. Says: "IV, Dryades - Dryas
Paphia, " The paphia of Hubner is considered to be paphia
Linnaeus, 1758, Since only paphia is included by Hubner,
it becomes the type of the genus.
Note, Hemming. 1958. Official Index Reject, and Invalid Works
Zool. Nomen, 1st, Install.: p, 4, no. 14, Hiibner's "Ten-
tamen" is included as a rejected work based on Opinion 97
of the Internat, Comm, Zool. Nomenclat,
2('3;. •229-259, 196}
ARGYNNID NAMES
233
Note, The name Dry as was later validly published by Hubner and
J?* E* Fabricius was selected by Hemming as its
type species.
MESODRYAS REUSS. 25 Jan. 1927. Deutsch. Ent. Zeitschr. (Berliner
Ent, Zeit, und Deutsche Ent, Zeit, Wieder, ), Jahrg. 1926 (5):
434-435 (in part). Proposes a new name for Dryas Hubner
[1806]^ as used in the "Tentamen" (which has been ruled in-
valid by the Internat, Comm, Zool, Nomen, ).
Type. P[apilio]. N[ymphalis]. [Phaleratus] paphia Linnaeus. 1758.
Syst. Nat. Toth. Ed. 1; 481, no. 138.
Type Selection. Reuss, As above. Says: ". . . Mesodryas T. Rss, ,
Type paphia L. , . "
ARGYREUS SCOPOLI. 1777. Introd. Hist. Natur. ; pp. 431-432, no.
171. Includes among others no. 208 (= P, N. [ P, ] niphe Lin-
naeus, 1767,
Type. P[apilio]. N[ymphalis]. [Phaleratus] niphe Linnaeus. 1767.
Syst. Nat. 12th, Ed. 1(2): 785, no. 208. (= Papilio N. hyper-
bius Johansson [or Linnaeus?], 1763 [after 23 Jixne]"^. Amoe-
nitates Academicae 6: 408-409, no. 7 5).
Type Selection. Reuss. 8 July 1928. Internat. Ent. Zeitschr. 22
(14): 146, Says of Argyreus: "...Die erstgenannte Art ist
hyperbius L, (= niphe L. , Syst, Nat, , ed. xii. Nr, 208) =
Type'5<f5i«y. " ” ^
Note. Linnaeus proposed niphe as a new name for hyperbius for
some undetermined reason.
Note, Hemming, 1958. Official List of Generic Names in Zoology.
1st. Install.: p. 101. no. 758, Argyreus is included on Of-
ficial List but does not have preference over Argynnis Fab-
ricius.
AC IDA LI A HUBNER. [1819]^. Verz. Bekannt. Schmett. (2): 31, no. 3.
Includes "A[cidalia], Niphe Linn. Syst, 308..."
Type. P[apilio]. N[ymphalis]. [Phaleratus] niphe Linnaeus. 1767.
Syst. Nat, 12th. Ed. 1 (2): 785, no. 208. (= Papilio N. hyper-
bius Johansson [or Linnaeus ?], 1763 [after 23 June]^^ Amoe-
nitates Academicae 6: 408-409, no, 75),
Type Selection. Scudder. 187 5 [Apr. ]^. Proc, American Acad. Arts
and Sci, 10 (art. v): 101, no, 12, Says of Acidalia: "Niphe may
therefore be taken as the type. "
Note. Linnaeus proposed niphe as a new name for hyperbius for
some xindetermined reason.
MIMARGYRA REUSS. Apr. 1922. Archiv fur Naturgesch. 87a (11):
221-222. Considers Mimargyra as a subgenus of Dryas as
used by Hubner in his "Tentamen" [1806].
Type, Papilio N[ymphalis]. hyperbius Johansson [or Linnaeus?],
1763 [after 23 June] Amoenitates Academicae 6: 408-409,
no. 7 5.
Type Selection. Reuss, As above. Says: "Mimargyra m. , subgen.
nov. , Type; hyperbius Joh, . , "
Note, Linnaeus proposed P. N. [P. ] niphe (1767, Syst. Nat, 1 (2):
785, no. 208) as a new name for hyperbius,
ARGYRONOME HUBNER. [1819]^. Verz. Bekannt. Schmett. (2): 32,
no. 6, Includes among others "265. A[ rgyronome], Laodice Pall.
Iter. 1. 61. "
234
McHENRY
/. Res. Lepid.
Type. Papilio Nymph[alis]. Phaleratus laodice Pallas, 1771, Reise
Verch. Provinzen Russischen Reichs 1: 470, no. 61,
Type Selection, Scudder. 1875 [Apr, ]5, Proc. American Acad, Arts
and Sci. 10(art. v); 120, no. 120, Says of Argyronome: "Should
Laodice prove generically distinct from the species of the
Dryas and Argynnis, this name may be reserved for it, , , "
Note. Fruhstorfer. [14May 1912]^^. In Seitz, Die Gross-Schmetter,
der Erde. 2nd. Abteil. 9(127): 514, Spells name as Argyro-
noma,
EUDAPHNE REUSS, Apr. 1922. Archiv fiir Naturgesch. 87a(l): 220-221
(in part). Considers Eudaphne as a subgenus of Dryas as used
by Hubner in his "Tentamen" of [1806]^,
Type, Papilio Nymph[alis]. Phaleratus laodice Pallas. 1771. Reise
Verch. Provinzen Russischen Reichs 1: 470, no, 61.
Type Selection, Reuss. As above. Says: "Eudaphne, m, subgen, nov. ,
Type: laodice Pall. . . "
BOLORIA MOORE. 10 Apr. 1900. Lepidoptera Indica 4(pp. 177-248):
243-244 (and pp. 245-254); plates 375-378).
Type. P[apilio]. pales [Denis and Schiffermilller]. 1775 [8 Dec, ]^.
Ankund, Syst, Werkes Schmett, Wienergegend; p. 177, no, 8
of K (pp. 176-178).
Type Selection, Moore. As above. Says of Boloria: "Type, - B,
Pales, Schiff, "
BRENTHIS HUBNER, [1819]^. Verz. Bekannt. Schmett., (2): 30, no. 2.
Includes among others "234. Brenthis Hecate Schiff. Verz, L.
4^ M ^
Type. P[apilio]. hecate [ Denis and Schiffermuller], 1775 [8 Dec.
Ankund. Syst, Werkes Schmett, Wienergegend; p, 179, no. 4
of L (pp. 178-179).
Type Selection, Scudder, 1872 [ J\ine]^„ Syst, Rev. Some Amer,
Butt, , pp, 24-25, Says of Brenthis: "Type Papilio Hecate
Wien, Verz. "
CHILDRENA HEMMING. 15 Feb. 1943, Proc, Royal Ent. Soc, London.
Ser, B. 12 (2): 30, Proposes a new name for Eudryas Reuss
which is preoccupied.
Type, Argynnis childreni G, Gray. 1831 [Feb,] . In J. Gray, Zool,
Misc. 1 (1): 31,
Type Selection, Hemming. As above. Says of Childrena: "Type Argy-
nnis childreni Gray, 1831, "
EUDRYAS "reuss. ^Oct. 1926. Internat, Ent, Zeitschr. 20(28): 253.
Type, Argynnis childreni G, Gray, 1831 [Feb. ]'^, In J, Gray. Zool.
Misc. 1 (1): 31.
Type Selection. Reuss. As above. Says: ", , , die neue Gattung: Eu-
dryas T. Rss. , Typ childreni. " The name childreni is of
Gray as per a later Reuss reference,
Eudryas Reuss is a homonym of Eudryas Boisduval. [10 Apr,
1 836]^, Hist. Nat. Ins. Spec. Gen, Lepid, Plates (Livrs,
1-2): p, 4 (explanation of plates); plate 14 (10b), fig, 9. Hem-
ming proposed Childrena as a new name,
CLOSSIANA REUSS. 30 Oct, 19’20. Entomolog. Mitteilungen 9(10- 12):
192 (foot-note).
Type, P[apilio]. selene [Denis and Schiffermuller], 1775 [8 Dec. ]'^.
Ankund. Syst, Werkes Schmett, Wienergegend; p, 321, no, 11
of K (pp, 176-178).
2(3):229-239, 1963
ARGYNNID NAMES
235
Type Selection, Reuss, As above. Says: "Clossiana m. , Type:
Selene, " In a later reference he says "selene Schiff, "
DAMORA NORDMANN. 1851, Bull. Soc, Imper, Nat. Moscou 24 (pt, 2^
no, 4): 439-442; plate 11, figs, 1-2,
Type, Damora paulina Nordmann, As above. Pages 440-442; plate
11, figs, 1-2, (= Argynnis sagana Doubleday and Hewitson. [4
Oct, 1847]*^. Genera Diur, Lepid, 1(12): plate 21, fig, 1. Text
on page 175, no, 18 was published later.
Type Selection, Nordmann, As above. Gives only "Damora paulina",
a new species which is considered to be a synonym of Argynnis
sagana Doubleday and Hewitson,
FABRICIANA REUSS, 30 Oct. 1920. Entomolog, Mitteilungen, 9(10-12):
192 (foot-note).
Type. P[apilio], N[ymphalis]. [Phaleratus] niobe Linnaeus. 1758.
Syst, Nat. 10th, Ed. 1: 481-482, no, 143.
Type Selection. Reuss. As above. Says: "Fabriciana m, , Type:
niobe". The name niobe is of Linnaeus as per a later Reuss
reference.
Note. Hemming, 1958, Official List of Generic Names in Zoology.
1st, Install.: 172, no. 1244. Fabriciana is included on Official
List.
IS SORIA HUBNER. [1819]^ Verz. Bekannt. Schmett. (2): 31, no. 2.
Includes among others "251. I[ssoria]. Lathonia Linn. [1767]
Syst, Pap, 213, " whichisin turn P. N. [P, ] lathonia Linnaeus,
1758. “ ~ “
Type, P[apilio], N[ymphalis], [Phaleratus] lathonia Linnaeus, 1758.
Syst. Nat, 10th. Ed. 1: 481, no 141,
Type Selection. Scudder, 1875 [Apr. ]^. Proc. American Acad, Arts
and Sci, 10 (art, v): 198, no, 577, Says (in part) of Issoria:
"1850. Steph, , Gat, Brit. Lep, 14: uses it for Lathonia only.
In which he is followed by Kirby (1858), and this therefore be-
comes the type.
RATHORA MOORE. 10 Apr. 1900, Lepidoptera Indica 4 (pp, 177-248):
241 (and pp. 242-243; plate 375).
Type, P[apilio]. N[ymphalis]. [Phaleratus] lathonia Linnaeus, 1758.
Syst, Nat. 10th. Ed. 1: 481, no, 141.
Type Selection. Moore, As above. Says of Rathora: "Type, - R,
Lathonia, Linn. "
K U K E N T H A L I E L L A REUSS. 1 Apr, 1921, Societas Entomologica 36
(4): 16,
Argynnis gemmata Butler, Jan. 1881. Ann, and Mag, Nat.
Hist. Ser. 5. 7 (37): 32, no. 1; plate 4, fig, 1,
Type Selection, Reuss. As above. Says: ", , . and Kiikenthaliella m.
(= Boloria part. ), Type gemmata Btlr. "
MESOACIDALIA REUSS, 10 Apr, 1926, Deutsche Ent, Zeitschr, (Ber-
liner Ent. Zeit, xind Deutsche Ent. Zeit. Wieder, ), Jahrg.
1926 (1): 69, subgenus no. 2 of genus no. VI (Acidalia Hubner).
Type. P[apiIio]. ^ymphalis], [Phaleratus] aglaja Linnaeus. 1758.
Syst, Nat. 10th. 1: 481, no. 140. (= P. N. P. charlotta
Haworth. 1803 [in or after July]^^), (I have not seen this
reference of Haworth).
Type Selection, Reuss. As above. Indicates the type when he says:
")Ie2, Subg. Mesoacidalia T. Rss., aglaia L, "
236
McHENRY
/. Res. Lepid.
Note. Hemming, 14 Nov, 194Z. Proc, Ent. Soc, London, Ser. B.
11(11): 155-160. Shows that the oldest name available for what
Linnaeus called P. N, [P. ] aglaja (1758, Syst, Nat, 10th, Ed,
1: 481, no. 140) is P. N. P. charlotta Haworth since the name
aglaja is a homonym of P, H, aglaja Linnaeus (1758, Syst.
Nat. 10th. Ed. 1: 465, rm. 44).
NEOACIDALIA REUSS. 10 Apr. 1926. Deutsche Ent. Zeitschr. (Ber-
liner Ent, Zeit. und Deutsche Ent. Zeit, Wieder. ). Jahrg,
1926 (1): 69, gives as a synonym of genus no. VI (Acidalia
Hubner).
Type. P[apilio], N[ymphalis]. P[haleratus]. cybele Fabricius, 1775
[17 Apr. Syst. Ent.; p. 516, no. 311.
Type Selection. Reuss, As above. Says: "VI. Acidalia Hbn. , cybele
Cram, (nur Pfeilduftschuppen). " The "cybele Cram. " of
Reuss is considered to be cybele Fabricius as Fabricius is
the author of the name; Cramer never used the name.
PANDORIANA WARREN. Nov. 1942. Entomologist: 75(954): 245-246.
Type. Pap[ilio]. Nymph[ali3]. Phalerat[us]. maja Cramer. [1776]"*,
Uitland. Kapellen Voorkom. Drei Waereld-Deelen Asia, Af-
rica en America 1 (8); 153, The text; p. 35, no. B-C; and
figures B-C on plate 25 were published earlier without a com-
plete name. (= P[apilio]. pandora [Denis and Schiffermuller].
1775 [8 Dec, Ankixnd. Syst. Werkes Schmetter. Wiener-
gegend; p, 176, no. 1 of K (pp. 176-178).)
Type Selection. Warren. As above. Says: "Genus Pandoriana gen.
nov. , type Papilio maja Cramer (= pandora Schiffermuller)".
Note. Hemming. 1958. Official List of Generic Names in Zoology.
1st, Install.: 177, no. 1270. Pandoriana is included on Official
List,
Note, The maja of Cramer and pandora [Denis and Schiffermuller]
are considered to indicate the same species.
PROACIDALIA REUSS. 10 Apr. 1926. Deutsche Ent. Zeitschr. (Ber-
liner Ent. Zeit. imd Deutsche Ent. Zeit. Wieder. ). Jahrg.
1926 (1): 69, subgenus no. 3 of Genus VI (Acidalia Hubner).
Type. Argynnis clara Blanchard, 1844, In Jacquemont, Voy, dans
Inde 4: 20, no. 14; plate 2, figs. 2-3. (I have not seen this
work of Blanchard).
Type Selection. Reuss. As above. Says: "5(t3. Subg, Proacidalia T.
Rss. , clara Blanch. " Since Reuss gives only clara, it be-
comes the type.
PROCLOSSIANA REUSS. 10 Apr, 1926. DeutscheEnt. Zeitschr, (Ber-
liner Ent. Zeit. \and Deutsche Ent. Zeit. Wieder. ), Jahrg.
1926 (1): 69, subgenus no, 1 of genus no, V (Clossiana Reuss),
Type, [Papilio] aphirape Hubner, [24 Dec. 1 799]-[ 1 3 Apr, 1800]^ ,
Samm, Europaischer Schmetter. (Papiliones): plate 5, figs.
23-25. Text on page 8, no. 1 was published later. (= Pap,
Nimph. Phal. eunomia Esper. [ 1799] ^ Die Schmetterlinge.
Suppl, Theils 1 (?): 94, no. 212. Plate 110 (? publication
date) contains fig, 5 which is lalelled "N, Phaler, Etmomia. "),
Type Selection, Reuss, As above. Says: "*1. Subg. Proclossiana T,
Rss, , aphirape Hbn. "
Note, The name aphirape Hubner is considered to be a synonym of
eunomia Esper, Hubner in his aphirape text cites eunomia
as a synonym. Dates of publication: eunomia (text): [1799]^^;
e\inomia (figure): ?; aphirape (figure): [24 Dec, 1799]-[13 Apr.
1800]^ i; aphirape (text): [21 Nov, 1805]-[31 Dec. 1805]11.
2(}):229-2}9, 196}
ARGYNNID NAMES
237
P R O F A B R I C I A N A REUSS. 10 Apr, 1926, Deutsche Ent. Zeitschr,
(Berliner Ent, Zeit. und Deutsche Ent. Zeit. Wieder. ). Jahrg.
1926(1); 70; given as an unnumbered subgenus of genus no. VII
(Fabriciana Reuss),
Type. Argynnis jainadeva Moore. [Nov, 1864]^^. In Lang, Ent, Mon.
Mag, I (6): 131.
Type Selection. Reuss. As above. Says: "Subg. Profabriciana T.
Rss, , jainadeva Moore. " Since Reuss gave only jainadeva, it
becomes the type,
PROKUKENTHALIELLA REUSS, 25 Jan. 1927. Deutsche Ent, Zeit-
schr. (Berliner Ent. Zeit. und Deutsche Ent. Zeit, Wieder. ),
Jahrg, 1926 (5): 435.
Type. Argynnis excelsior Butler, 1 Apr. 1886. Proc. Zool, Soc,
London forT895 (4): 729-730, no. 42.
Type Selection. Reuss. As above. Says; ". . . Prokukenthaliella T.
Rss, , subg. nov. , Typ excelsior Butl. . . "
PROTODRYAS REUSS, 8 July 1928. Internat. Ent. Zeitschr. Guben
22(14): 146. Proposes Protodryas to replace Prodryas Reuss
which is a preoccupied name.
Type. Argynnis kamala Moore. 1857 [after 7 Dec.] , In Horsfield
and Moore. Catal, Lepid. Ins. Mus. Hon. East-India Co. 1:
156-157, no. 324.
Type Selection. Reuss. As above. Says; "...Protodryas T. Rss, gen.
nov, , Typ kamala Moore. . . "
PRODRYAS REUSS, 10 Apr. 1926. Deutsche Ent. Zeitschr. (Berliner
Ent, Zeit, und Deutsche Ent, Zeit, Wieder,), Jahrg, 1926 (1):
66; subgenus no. 1 of genus no. I (Dryas Hubner).
Type. Argynnis kamala Moore. 1857 [after 7 Dec, In Horsfield
and Moore. Catal. Lepid, Ins, Mus. Hon. East-India Co. 1:
156- 157, no. 324.
Type Selection, Reuss, As above. Says: "^til. Subg, Prodryas T.
Rss, , kamala Moore. "
Note, Prodryas Reuss is a homonym of Prodryas (fossil) Scudder,
3 May 1878. Bvdl. U. S. Geol. and Geogr, Surv. , 4(2): 520.
Reuss later proposed the name Protodryas to replace his Pro-
dryas.
PSEUDORA THORA REUSS. lOApr. 1926, Deutsche Ent. Zeitschr.
(Berliner Ent. Zeit. und Deutsche Ent, Zeit. Wieder.). Jahrg,
1926 (1): 68, subgenus no. 1 of genus no. III (Rathora Moore).
Type. Argynnis issaea (Nomen nudum) Doubleday (G. Gray Ms. name).
[2 June 1848]'^. In Doubleday and Hewitson, Genera Diur. Lepid,
1(20): 174, no. 3. (= Argynnis issaea Moore (G. Gray Ms,
name), 1857 [after 7 Dec. ] f 4:^ in Horsfield and Moore, Catal,
Lepid, Ins, Mus, Hon, East-India Co. 1; 156, no, 323),
Type Selection. Reuss, As above. Says: "*1. Subg. Pseudorathora
T. Rss, , isaeea Dbldy, , isaeoides T. Rss, Ztrlgb. " Note:
Reuss probably intended to nominate isaeea.' as the type species
but did not do so. If no type has subsequently been named, I
will here designate "isaeea Dbldy, ", used by Reuss, as the
type of the genus Pseudorathora Reuss but noting the identity
of the species in accord with the data presented in the following
note.
238
McHENRY
/. Res. LepiJ.
Note. The name Argynnis issaeahas an involved history. It was ap-
parently first used as a Nomen nudum by G, Gray in his
Lepid. Nepal, p. 11, 1846. It was later used as a Nomen
nudum (G. Gray Ms. name) by Doubleday in Genera Diur,
Lepid, 1: 174, no. 3^ Moore (in Horsfield and Moore, Lepid.
Ins. Mus. Hon. East-India Co, 1: 156, no, 323) would appear
to be the author since he applies the name to Kollar's de-
scription of lathonia Linnaeus (not Linnaeus) in Hugel's
Kaschm, 4: 440, 1848, Reuss misspelt the name as isaeea,
(I have not seen the works of Gray and Kollar),
S EMNOPYSCHE SCUDDER. 1875 [Feb. -6Mar.]15^ Bull. Buffalo
Soc. Nat. Sci. 2 (4) signs. 30-34: 238, no. 52; pp, 258-259,
no. 44.
4
Type. Pap[ilio]. Nymph[alis]. Phaler[atus] diana Cramer. [1777] ,
Uitland. Kapell. Voorkom, Drie Waereld-Deelen Asia, Af-
rica en America 2(16): 148. Described and figured earlier,
without a name, on page 4, figs, D-E and on plate 98, figs,
D-E in Vol. 2, pt. 9.
Type Selection. Scudder. As above. Says of Semnopysche: "Type:
Papilio Diana Cram. "
SPEYERIA SCUDDER. 1872 [June]^. Syst. Rev. Some Amer, Butt,,
P, 23.
Type, Nym[phalis]. Phal[ eratus], idalia Drury. [1773] . Illust, Nat.
Hist. 1: [1] in Index. Described and figured earlier (without a
name) on page 25, figs. 1-3 and plate 13, figs. 1-3,
Type Selection. Scudder. As above. Says of Speyeria: "Type Papilio
Idalia Drury. "
YRAMEA REUSS. 30 Oct, 1920. Entomolog. Mitteilxingen 9( 1 0- 1 2): 192
(foot note).
Type. Nym[phalis]. Ph[aleratus]. cytheris Drury. [1773]'^. Illust,
Nat. Hist. 2: 7, figs. 3-4; plate 4, figs. 3-4; p. [1] in Index,
Type Selection, Reuss. As above. Says: "^ ) Y ramea m. , nov. gen,.
Type: cytheris Drury (1773), "
CHILARGYNNIS BRYK. 6 Nov. 1944. Archiv fiir Zoologi 36a(3): 8-9
(9-10, in part). ^
Type, Nym[phalis]. Ph[aleratus]. cytheris Drury. [1773] . Illust.
Nat. Hist, 2: 7, figs. 3-4; plate 4, figs, 3-4; p, [1] in Index.
Type Selection. Bryk. As above. Says: "Genus Chilargynnis m, (gen.
nov,). Typus generis: Nymph. Ph, cytheris Drury, 1773."
FOOTNOTES
Vol. 6 title page date is qualified by indirect date data (certain
names in Fabricius' article were mentioned in the Allgemeine Liter-
atur-^Zeitung, Halle [Jena] 2(303): cols, 1177-1181, 19 Dec. 1807).
2. Hemming. 19 58. Official Index Reject, and Invalid Workes in Zool,
Nomencl. 1st. Install.: p. 4, no. 14.
3. Vol. 6 title page signature is qualified by article date,
4. Hemming. 1958. Official List Works Approv. Avail, Zool. Nomencl,
1st, Install, : p. 4, no. 15 (dates for Hubner's Verz. Bek. Schmett, );
p. 7, no. 30 (dates for Vols, 1-3, Drury's Illust. Nat, Hist, ); p. 9,
no. 34 (date for Fabricius ' Syst. Ent. ); p. 9, no. 37 (date for [Denis '
and Schiffermuller ' s] Ankilnd, Syst. Werkes Schmett. Wienergegend);
pp. 9-10, no. 38 (dates for Cramer's and Stoll's Uitlandsche Kap-
pelen).
-229-259, 1965
ARGYNNID NAMES
239
5. Vol. 10 title page signature date is qualified by date of receipt given
for article in Psyche, Jour. Ent. 2(33-34); 9, no. 716, 1883.
5a. Griffin. 1936, Trans. R. Ent. Soc. Lond. 85: 243-279.
6. V/ork title page date is qualified by date of a review in June 1872 num-
ber of Amer. Nat. 6: 354-359.
7. Sherborn. 1922. Index Animalium, 2nd. Ser. Vol. A-B: cxxxi.
8. Scudder. 1899. Psyche, Jour. Ent. 8: 153- 154.
9. Hemming,' 1936- 1943. Jour, Soc. Bibliog. Nat. Hist. 1: 335-464.
10. Work title page signature date is qualified by the preface date (p.
xxx).
11. Hemming. 1937. Hilbner 1: 146-324.
12. Sherborn and Woodward. 1901. Ann. and Mag. Nat. Hist. Ser, 7.
7: 137-140.
13. I have not seen the wrappers for Vol. 1 which apparently gave the
dates and numbers for each respective number. On the basis of the
plan for later volumes I have established this date.
14. Vol. 1 title page date is qualified by the preface date (p. v).
15. Vol. 2 title page signature date is qualified by the date of signatures
30-34 and the date of receipt of a copy of the article (ex Cambridge
Ent. Club, copy now in my library).
LIST OF REFERENCES
Dos Passos, Cyril Franklin and Lionel Paul Grey.
1945. A genitalic survey of Argynninae (Lepidoptera, Nymphalidae).
American Mus. Novitates (1296): 1-29; figs. 1-54.
Doubleday, Edward.
1848. In Doubleday, Hewitson and Westwood (in part). The Genera of
Diurnal Lepidoptera. .. 1 ; 168- 170 (Euptioeta).
Linnaeus, Carolus.
1758. Syst. Nat. 10th. Ed. 1: 458-488, no. 203 (Papilio).
Reuss, F. A. Theodore.
1926. Systematischer Uberblick der Dryadinae T. Rss. , mit einigen
Neubeschreibungen. Deutsche Ent. Zeitschr. Jahrg. 1926:
65-70.
[1927]. Uber Funktion der Sexualarmaturen bei Lepidopteren (Rhop. )
und die r esultierende Weiter entwicklung meines versuchten
natiirlichen Systems der Dryadinae T. R. Ibid. Jahrg, 1926:
431-440. ~
Warren, Brisbane Charles Somerville.
1944. Review of the classification of the Argynnidi: with a systematic
revision of the genus Boloria (Lepidoptera; Nymphalidae).
Trans. R, Ent. Soc, Lond. 94: 1-153; plates 1-68; text figs.
1-14.
1955. A review of the classification of the Subfamily Argynninae
(Lepidoptera: Nymphalidae). Part 2. Definition of the Asiatic
genera. Ibid. 105; 381- 391; plates 1-4, each with an explanation
page.
Warren, Brisbane C. S. , Cyril F. Dos Passos and Lionel P. Grey.
1946. Supplementary notes on the classification of Argynninae (Lepid-
optera: Nymphalidae). Proc. R. Ent. Soc. Lond. (B) 15: 71-73.
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YOSEMITE BUTTERFLIES
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THE LEPIJDOFTER^A
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Number 3
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IN THIS ISSUE
Affinities and distribution of Antillean Ithomiidae
Richard M. Fox 173
Ovipositional preference tests with Pieris
William Hovanitz and Vincent C.S. Chang 185
Notes on the early stages of Drepanulatrix monicaria
(Guenee) ( Geometridae ) John Adams Comstock 201
Biographical sketch — John Adams Comstock 204
The origin of a sympatric species in Colias through the aid
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Biographical sketch — James Wilson Tilden 224
A rubber stamp method for producing specimen labels
Phillip A. Adams 225
Generic or subgeneric names closely related to Argynnis
Paddy McHenry 229
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Volume 2 Number 4 December, 1963
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© Copyright 1964
EVIDENCE EOR LACK OF TERRITORIALITY IN
TWO SPECIES OF HAMADRYAS ( NYMPHALIDAE )
GARY N. ROSS
Louisiana State University, Dept, of Entomology,
Baton Rouge, Louisiana
A TERRITORY AS DEFINED BY Kendeigh' (1961) “is any area
defended against intruders.” Today, it is a well known fact that
many animals have well established territories for either their
entire lives or else for only brief periods during their lives, e.g.,
during the breeding season or at feeding times. This behavior
pattern frequently has been reported for vertebrates — birds
(Hinde, 1956), mammals (Seton, 1909, 1925-28), amphibians
(Sexton, I960), fish (Hasler and Wisby, 1958), turtles (Cagle,
1944 ) ; some invertebrates — snails ( Edelstam and Palmer, 1950 ) ,
wood ants (Elton, 1932), dragonflies (Jacobs, 1955). In regards
to butterflies, it is known that many species (Phyciodes tharos,
Precis coenia, Lycaena phLeas, Danaus plexippus, several angle
wings of the genus Polygonia) are pugnacious, chasing any
moving object that gets within close range ( Klots, 1951 ) . It even
has been recorded (Klots, 1951) that males of Lethe sp. adopt
favorite perches on tree trunks which they resume day after day,
darting out occasionally in order to drive away other males.
If this is the case, it can be said that these species exhibit ter-
ritoriality. However, there is no experimental evidence to sup-
port this idea which is based solely on field observations.
Closely related to territoriality, is the subject of home range —
“that area regularly traversed by an individual in search of food,
mates and caring for young” (Kendeigh, 1961). The few ex-
periments undertaken with butterflies (Dowdeswell, Fisher and
Ford, 1940, 1949; Evans, 1955; Abbott, 1959; Faler, 1959; Ehrlich,
1961) show that some species of butterflies do not wander very
much during the course of their lives but rather are sedentary
and tend to remain in relatively small areas. In other words,
they have small home ranges.
The present paper sheds a little light on these two areas of
behavior regarding two nymphalids, Hamadryas februa gudula
Fruhstorfer and Hamadryas guatamalena guatamaleria Bates.
241
242
ROSS
/. Res. Lepid.
While engaged in collecting butterflies in the Tuxtla Moun-
tains, Veracruz, Mexico (summer, 1962), I became aware that
there were fairly large populations of both Ha7nadryas februa
and Hamadnjas guatamalena in a small ( two acres ) tract of land
given over to the cultivation of coffee. These buttei*flies were a
conspicuous part of the landscape for whenever any moving
object entered the immediate area, individuals which previously
were resting on the trees, darted out, making the peculiar click-
ing noises that are so characteristic for the group (Ehrlich &
Ehrlich, 1961). After a few seconds of rapid zig-zag motions,
they again usually took up resting positions on either the same
trees or else on others in the same general vicinity.
The attitudes of rest for these two species are quite interest-
ing. Individuals rest head downward on the trunk or larger
limbs with their wings stretched out horizontally and held flat
against the bark. In this position, individuals are very much in-
discernable for the somber color patterns of the dorsal side of
the wings blend in perfectly with the bark and its lichen en-
crustations. Trees of the species Inga spuria Humb. & Bonpl.
( Leguminosae ) which has been planted for their value as shade
trees in coffee culture were the tallest in the area and were most
often used for resting.
After collecting in this area for several days, I noticed that
many of the 7. spuria trees habitually seemed to have one or
more individuals of 77. februa and 77. guatamalena resting on
them, A good friend of mine (and an ardent butterfly collector,
also) remarked that he had observed this same type of phe-
nomenon throughout Mexico and parts of Central America and
that he thought that individuals rested repeatedly on certain
apparently “favored” trees and defended these positions against
all intruders. If this were the case, both species exhibited ter-
ritoriality. This had to be determined.
In this same tract were numerous orange trees. These (the
larger trees in particular) served as excellent feeding stations
for both species of Hamadryas as well as for several species of
Prepona, Smyrna, Anaea and Caligo. Termites being numerous
in the area, frequently made excavations into the bark and sap
wood of these trees and consequently caused sap to ooze out
from the burrowings. This sap combined with moisture (either
from heavy morning dew or from rainwater) gave off a strong
acrid odor which was quite noticeable to me for a distance of
a meter or more. Apparently this fermenting sap was highly
2 (4):241-246, 1963
TERRITORIALITY
243
prized by the above mentioned butterflies for at such times ( when
conditions were favorable for production of good quantities of
sap and for the fermentation thereof), large numbers of them
were seen feeding on the material. Because of the high pop-
ulation densities of Hamadryas in the area, it was a common
everyday sight to -see several individuals of each species feeding
simultaneously on these trees. When a new arrival tried to join
the feeding association, it usually was chased away. However,
sometimes it was able to force its way into the group and thus
take up a feeding position. It had to be determined whether
territorialism was being exhibited here, that is, whether or not
the same individuals were feeding at the stations day after day.
In an attempt to get some quantitative data concerning the
above phenomena, three simple capture-recapture experiments
were performed.
Experiment I: Resting habits of Hamadryas februa gudula
Fruhst.
Three trees which, after several daily inspections proved to
be fairly constant resting places for H. februa, were selected and
marked, each with a distinct and different color. The trees were
situated in a triangular pattern and were approximately 10 meters
apart. All butterflies on these trees were captured, marked
using the l-2~4-7 system of Ehrlich & Davidson (1960), and
immediately released. At the same hour on the following day,
the trees were inspected and all butterflies found resting upon
them were captured, checked and released. Any new, unmarked
individuals were marked. All other trees within a distance of
20 meters in all directions were checked, also. It should be
mentioned here that when approaching the trees, considerable
caution was exercised so that the butterflies were caused the
least amount of disturbance. This same procedure was followed
for 11 consecutive days.
The results of this experiment proved to be rather interesting.
Of 24 individuals ( 20 ^ 5,4? $ ) of H. februa marked during the
11 days of the experiment, only 5 specimens were recaptured
on the same trees from which they were taken originally. Of
these 5, only 3 were captured more than once on the original
tree. Of these same 24 individuals, only 9 were recaptured on
nearby trees. Thus, it appears that most individuals do not
regularly frequent the same trees or even other similar trees
in the nearby area, and that there is a constant influx of new
individuals from other surrounding areas.
244
ROSS
J. Res. Lepid.
Experiment II: Resting Habits of Hamadryas guatamalena
GUATAMALENA BaTES
Two 1. spuria trees which seemed to be visited frequently by
members of H. guatamalena were selected. All butterflies found
resting on them were captured, marked and released. The
following day the trees were inspected and all individuals found
upon them were checked. All new, unmarked specimens were
marked and released. Again, all other trees within a distance of
approximately 20 yards were checked. This procedure con-
tinued for 7 consecutive days.
Of 9 individuals ( 7 ^ $ y 29 9 ) of H. guatamalena marked,
none were recaptured on the trees from which they were taken
originally. Of these 9, only 2 were recaptured in the immediate
vicinity. Thus, it appears that H. guatamalena also does not
habitually frequent the same resting spots but that new in-
dividuals enter the area from adjacent habitats.
Experiment III: Feeding Habits of H. februa and
H. GUATAMALENA.
As mentioned previously, several of the Citrus trees in the
study area served as excellent feeding stations for both species
of Hamadryas. In an attempt to determine if feeding territories
existed for the species, two highly productive trees situated
about 10 meters apart were marked with different colors and all
specimens of Hamadryas found on them were marked with
corresponding colors. The same procedure of checking was
followed here.
After 7 consecutive days of checking, the following results
were obtained. Of a total of 15 individuals ( 5 ^ S , 39 9 oi H.
februa and 3 ^ S y 49 9 oi H. guatamalena) marked, none were
recaptured on the marked trees or on any other trees in the
general vicinity. Thus, it appears that no evidence exists for the
presence of feeding territories in either species.
DISCUSSION
It appears from the three simple capture-recapture studies
described above, that there is no evidence for territoriality in
either Hamadryas februa or H. guatamalena, two species of
nymphalids which upon first observations, appear to be rather
sedentary in their habits; it appears that these species do not
frequent the same rest spots daily nor do they have preferred
feeing places. From field observations, it appears that the
resting spots are gotten on a “first come, first serve’' basis and
that the more aggressive individuals usually gain control of the
desired spots. These same individuals in turn retain their
2 (4):241-246, 1963
TERRITORIALITY
245
positions until driven off by intruders or until intrinsically
motivated to go elsewhere. Therefore, I think that it is better
to say that these 2 species are very pugnacious.
One might argue that since these positions are defended,
their exists at least a sort of “transient territoriality.” However,
close field observations do not bear this out; after the departure
of an intruder, individuals may or may not return to the same
spots. Indeed, it appears that there is an equal chance of them
selecting other resting areas. Therefore, I do not believe that
one can say that these species have even “transient territories.”
It is interesting to speculate on the “whereabouts” of the
marked individuals. Only on one occasion was a marked
specimen recovered outside of the study area. This recovery
was made 6 days subsequent to the termination of the three
experiments. The insect, a male specimen of H. februa, was
feeding on the exuding sap of an orange tree which was growing
in a cow pasture approximately 100 yards from the original study
area. It seems improbable that the marked individuals were in-
jured during the actual marking process for no evidence for this
was observed. That this marking technique can be employed
with even smaller species of nymphalids without any apparent
ill-effects to the specimens, was shown by Ehrlich & Davidson
(1960) in their work with the checkerspot Euphydryas editha
Boisduval. To be sure, the question of predation does arise
here: were the marked individuals more susceptible to attacks
by such things as lizards or birds? I believe that this question
can be answered in the negative for as mentioned above, in-
dividuals of both species do not fly a great deal but spend a
large amount of their time resting with their wings expanded
horizontally and held flat against the bark. In such positions,
the small artificial marks on the undersurfaces are completely
invisible. Furthermore, flights when made, are of short duration
and of such a rapid nature as to cause little chance of the mark-
ings becoming detected by predators. Therefore, I think that
it is highly probable that marked individuals simply wandered
off to surrounding areas and became widely dispersed amongst
the multitude of other individuals. I think, then, that it is safe
to conclude that these two species do not have small home
ranges. An interesting future experiment would be to enlarge
the study area and to determine the actual dispersion distances.
In conclusion, it seems that there is no evidence for ter-
ritoriality in either Hamadryas februa or Hamadryas guata-
malerm.
246
ROSS
/. Res. Lepid.
ACKNOWLEDGEMENTS
I would like to express my appreciation to the following
people: Dr. F. M. Brown (Fountain Valley School, Colorado
Springs, Colorado) for butterfly determinations; Dr. V. E. Rudd
(U. S. National Museum, Smithsonian Institution, Washington,
D. C. ) for plant determinations; Mr. S. L. Warter, Dr. M. S.
Blum, and Dr. L. D. Newsom (Louisiana State University,
Entomology Research Department) for criticisms regarding the
manuscript; Mr. and Mrs. R. F. Andrle (Buffalo, New York) for
living accommodations; Christopher Andrle (Buffalo, New York)
for his assistance with the field work.
LITERATURE CITED
ABBOTT, W. 1959. Local autecology and behavior in Pieris protodice
Boisduval & Leconte with some comparisons to Colias eunjtheme
Boisduval ( Lepidoptera: Pieridae). Was. J. Biol. 17: 279-298.
CAGLE, F. R. 1944. Home range, homing behavior, and migration in
turtles. Misc. Puhl. Mus. ZooL, Univ. Michigan, 61: 1-34.
DOWDESWELL, W. H., R. A. FISHER, & E. B. FORD. 1940. The
quantitative study of populations in the lepidoptera. 1. Polyomniatus
icarus Rott. Ann. Eugenics 10: 123-136.
1949. The quantitative study of populations in the lepidoptera.
II. Maniola jurtina L. Heredity 3: 67-84.
EDELSTAM, C. and C. PALMER. 1950. Homing behavior in gastro-
podes. Oikos 2: 259-270.
EHRLICH, P. 1961. Intrinsic barriers to dispersal in eheckerspot butterfly.
Science, 134, no. 3472: 108-109.
EHRLICH, P. & A. H. 1961. How to know the butterflies. Wm. C. Brown
Pub. Co., Dubuque, Iowa, 262 pp.
EHRLICH, P. R. & S. E. DAVIDSON, 1960. Techniques for capture-
recapture studies of lepidoptera populations. Journ. Lep. Soc. 14:
227-229.
ELTON, C. 1932. Territory among wood ants (Formica rufa L.) at
Picket Hill. J. An. Ecol. 1: 69-76.
EVANS, W. H. 1955. Retrieving marked Anthocaris reakertii. Lep. News
9:118.
FALES, J. H. 1959. A field study of the flight behavior of the Tiger
Swallowtail Butterfly. Ann. Ent. Soc. Am. 52: 486-487.
HASLER, A. D. and W. J. WISBY. 1958. The return of displaced large-
mouth bass and green sunfish to a “home” area. Ecol. 39: 289-293.
HINDE, R. A. 1956. The biological significance of tlie territories of birds.
Ibis 98: 340-369.
JACOBS, M. E. 1955. Studies on territorialism and sexual selection in
dragonflies. Ecol. 36; 566-586.
KENDEIGH, S. C. 1961. Animal ecology. Prentice-Hall, Inc., Engle-
wood Cliffs, N. J., 468 pp.
KLOTS, A. B. 1951. A field guide to the butterflies of North America,
east of the Great Plains. Houghton Mifflin Company, Boston, Mass.,
349 pp.
SETON, E. T. 1909. Life histories of northern animals. Charles Scribner’s
Sons, New York, N. Y., 2 vols.
1925-28. Lives of game animals. Doubleday, Doran Col, New
York, N. Y., 4 vols.
SEXTON, O. J. 1960. Some aspects of the behavior and of the territory
of a dendrobatid frog, Prostherapis trintatis. Ecol. 41 (1): 107-115.
Journal of Research on the Lepidoptera
2(4) :247-270, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 1964
A SYNOPSIS OF THE WEST INDIAN LYCAENIDAE,
WITH REMARKS ON THEIR ZOOGEOGRAPHY
HARRY K. CLENCH
Carnegie Museum, Pittsburgh 13, Penna.
The basic reference for students of the Antillean Lycaenidae
is the thorough revision of Comstock & Huntington (1943). The
particular strength of this work is in the detail at species and subspecies
level, in the very large amount of material so profitably studied that
few major subsequent distributional discoveries have been or are likely
to be made, and in the meticulous and extremely perceptive study of
the older literature that underlies the whole work, resulting in a sta-
bility of species nomenclature that many a better known area might
well envy.
Problems of higher classification in the family, however, were almost
completely passed over. Reason for this is not hard to discover: the
problems involved so far transcended the boundaries of the West Indies
that to resolve them would have necessitated an effort out of all
proportion to the study and very likely would have greatly postponed
or even completely stopped its appearance.
Subsequent work by others now has made considerable progress in
reducing that hiatus. Of particular significance is the revision of some
of the blues by Nabokov (1945), who brought an intimate knowl-
edge of palearctic as well as nearctic groups to bear on the neotropical
fauna, where this knowledge was peculiarly valuable. Much later
my own work on the reclassification of the North American hairstreaks
(Clench, 1961) touched on many West Indian groups and indicated
that the male genitalia in particular could be of major assistance in
determining their relationships.
Some time ago Mr. F. Martin Brown asked if I would go over the
Jamaican hairstreaks for him in order to bring their generic classifica-
tion into line with the North American revision. This took more time
than at first expected, particularly in the collateral examination of
mainland species, but when the task was done it was evident that a
very little additional work would extend the results from just one
island to the whole West Indian region. In view of the many changes
indicated by Nabokov’s work and of an unpublished survey of my
own of Leptotes (world-wide), prepared by a fortunate coincidence
for quite another purpose, it seemed opportune to combine all this
into a synoptic list, herewith presented (Table I).
247
248
CLENCH
/. Res. Lepid.
TABLE 1
SYNOPSIS OF THE WEST INDIAN LYCAENIDAE
family LYCAENIDAE
subfamily STRY MONINAE
tribe EUMAEINI
Eumaeus Hubnerl819
L a tala Poey 183Z (Cuba, Bahamasu)
a. florida Rober 19 26(Florida)
=g r a y i Comstock and Huntington 1943
tribe STRYMONINI
Chlorostrymon Clench 1961
Z, maesites Herrich-SchJiffer (Florida, Bahamas,
Cuba, Jamaica, Puerto Rico^ not yet taken on
Hispaniola but should occur])
a. clenchi Comstock and Huntington 1943 (Dominica)
3. simaethis Drury 1770 (Cuba, Hispaniola, Puerto
Rico, Virgin Islands, St. Kitts, Dominica, St. Vincent)
a. jago Comstock and Huntington 1943 (Jamaica)
Othei subspecies in Central and South America
Calycopis Scudder 1876
4. be on Cramer 1780 (Hispaniola), Status uncertain
Nesiostrymon new genus (see note no. 1 )
5. cell da Lucas 1957 (Cuba)
a. shoumatoffi Comstock and Huntington 1943 (Jamaica)
b. aibonito Comstock and Huntington 1943 (Puerto Rico)
[not recorded from Hispaniola but probably occurs there]
Callophrys Billberg 1820 ("see note no. 2")
subgenus Cyanophrys Clench 1961
6. crethona Hewitson 187 3 (Jamaica)
Ps eudolycaena Wallengr en 1859 (see note no. 3)
7. marsyas Linnaeus 17 58 (South America)
a, cybele Godman and Salvin 1896 (St, Vincent)
Another subspecies in South America
Heterosmaitia new genus (see note no, 4)
8. bourkei Kaye 1924 (Jamaica)
Allosmaitia new genus (see note no, 5)
9. coelebs Herrich-Schaffer 1862 (Cuba)
10. fidena Hewitson 1867 (Hispaniola, Puerto Rico)
• 11. piplea Godman and Salvin 1896 (St. Vincent, Grenada,
Dominica)
Strymon Hibner 1818
12. a c i 8 Drury 1770 (Antigua Dominica*)
a. mars Fabricius 1777 (Puerto Rico,*Virgin Is, , St. Kitts)
b. petioni Comstock and Huntington 1943 (Hispaniola)
c. casasi Comstock and Huntingto n (Cuba)
Carpenter and Lewis (1943) report two subspecies from th^ Cayman Is,,
one from Grand Cayman and one from Little Cayman and Cayman Brae,
The identity of neither is known to me,
d. armouri Clench 1943 (Bahamas)
e. subspecies ? (Chicos Islands: Rindge 1955)
2 {4) 1247-270, 196}
WEST INDIAN LYCAENIDAE
249
f. g o s 8 e i Comstock and Huntington 1943 (Jamaica)
g. bartrami Comstock and Huntington 1943 (Florida)
13. martialis Herrich-Schaffer 1864 (Florida; Bahamas;
Cuba; Jamaica; Little Cayman, Cayman Islands)
14. rufofusca Hewitson 1877 (St. Vincent; [Central
America])
15. bubastus Cramer 1780 [South America, including
Trinidad]
a. ponce Comstock and Hintington 1943 (Puerto Rico;
Virgin Islands; St. Kitts; Antigua; Dominica; St. Lucia;
St, Vincent; Grenada)
16. columella Fabricius 1793 (Hispaniola)
a. modesta Maynard 1873 (Florida; Dry Tortugas *)
b. cybira Hewitson 1874 (Jamaica; Cuba; Bahamas*^
Grand Cayman, Cayman Islands ji)
c. arecibo Comstock and Huntington 1943 (Puerto Rico)
Other subspecies in Central America and related islands.
* 17. antigua Comstock and Huntington 1943 (Antigtia,
Virgin Islands)
*18. toussainti Comstock and Huntington 1943 (Hispaniola)
* 19. cristophei Comstock and Huntington 1943 (Hispaniola)
* 20. limenia Hewitson 1868 (Cuba; Jamaica; Hispaniola;
Puerto Rico)
21, bazochii Godart 1822 [Central and South America]
a, g un d 1 a c h i a n u s Bates 1935 (Cuba; Jamaica;
Hispaniola)
Electrostrymon Clench 1961 (see note no. 6)
22. pan Drury 177 3. (Jamaica)
23. angelia Hewitson 1874 (Cuba)
a, pantoni Comstock and Huntington 1943 (Jamaica)
b, dowi Clench 1941 (Bahamas)
c, boyeri Comstock and Huntington 1943 (Hispaniola;
Puerto Rico; Virgin Islands -St. Croix, 4, v. 1941,
leg. Beatty: 2 males, Carnegie Museum)
24, angerona Godman and Salvin 1896 (St, Kitts; St. Vincent;
Dominica)
* 25. dominicana Lathy 1904 (Dominica)
subfamily PLEBEJINAE
tribe CATOCHRYSOPINI
Leptotes Scudder 1876 (see note no. 7)
(a) c a 8 s i u s group
26. cassius Cramer 1775 ([South America]; possibly
Martinique)
a. catilina Fabricius 1793 (Virgin Islands; St. Kitts)
b. theonus Lucas 1857 (Florida; Bahamas; Cuba; Grand
Cayman, Little Cayman and Cayman Brae, Cayman
lslands;<0^amaica; Hispaniola; Puerto Rico)
c. Chadwick i Comstock and Huntington 1943 (Dominica;
Gaudeloupe)
Other subspecies in Central America,
(b) marina group
?27. marina Reakirt 1868 (Puerto Rico, probably as casual
or misidentified t h e o n u s; [Central America] )
28. perkinsae Kaye 1931 (Jamaica)
250
CLENCH
/. Res. Lepid.
tribe PL EBEJINI (see note no. 8)
Hemiargus Hffbner 1818
subgenus Hemiargus Httbner 1818
29. ceraunus Fabricius 1793 (Jamaica; Hispaniola;
Navassa)
a. filenus Poey 1832 (Cuba; Bahamas; Grand Cayman,
Cayman Islands *)
b. antibubastus Httbner 1818 (Florida; Georgia; etc.)
Other subspecies in Central America, south to Costa Rica.
30. hanno Stoll 1790 [South America, north to Costa Rica]
a. watsoni Comstock and Huntington 1943 (Hispaniola;
Puerto Rico; Virgin Islands; St. Kitts; Antigua;
Guadeloupe; Dominica; Martinique; St. Lucia; Grenada;
Barbados)
Other subspecies in South America,
subgenus Echinargus Nabokov 1945
31. isola Reakirt 1866 (Puerto Rico, probably casxial;
[Mexico])
Other subspecies on the mainland, from western United
States to Costa Rica.
subgenus C y c 1 a r g u s Nabokov 1945 (see note no. 9)
32. dominie a Moschler 1886 (Jamaica)
33. a mm on Lucas 1857 (Cuba; Bahamas; Grand Cayman,
Cayman Islands)
34. woodruff! Comstock and Huntington 1943 (Puerto Rico;
Virgin Islands; St, Kitts)
35. er embis Nabokov 1948 (Little Cayman Island, Cayman
Brae, Cayman Islands)
36. thomasi Clench 1941 (Bahamas)
a. bahamensis Clench 1943 (Crooked Island, Bahamas,
where it appears to be constant)
b. bethune-bakeri Comstock and Huntington 1943
(Florida)
c. noeli Comstock and Huntington 1943 (Hispaniola; Mona
Island)
Pseudochrysops Nabokov 1945
37. bornoi Comstock and Huntington 1943 (Hispaniola;
Puerto Rico [casual?] )
subfamily BREPHIDIINAE
Brephidium Scudder 187 6 (see note no, 10)
38. exilis Boisduval 1852 [western United States through
Central America]
a. isophthalma Herrich-Schaffer 1862 (Bahamas;
Cuba; Hispaniola; Jarraica; Grand Cayman, Cayman
Islands)
=thomp«soni Carpenter and Lewis 1943
* 39. barbouri Clench 1943 (Bahamas) Status uncertain
40. pseudofea Morrison 1873 (Florida; Georgia; northern
Bahamas)
2 {4) -.247-270, 1963
WEST INDIAN LYCAENIDAE
251
Ail taxonomic changes and additions are provided in a series of
notes referring to this list (Table I). The format of the list
needs but few remarks. References are not given but may be obtained
readily from Comstock & Huntington (1943) or some other work
listed at the end. Distributions are complete so far as I know them.
Species with an asterisk preceding their number have not been exam-
ined genitalically;' all others have. An asterisk immediately following
a locality indicates that the taxonomic status of the form in question
is uncertain on the island.
Bates (1935: 188) remarked concerning the West Indian Lycaen-
idae, "The family at present, then, is of almost no use to . . . the zoo-
geographer," a remark which was certainly true enough when it was
written. Comstock & Huntington’s work made a considerable improve-
ment, but left continental relationships, as before, quite uncertain.
Nabokov’s work and the present reclassification of the hairstreaks have
both paid particular attention to these continental affinities, and as a
result there are few forms whose relationship to mainland species or
groups is still doubtful. The time, in other words, has come for a
zoogeographic assessment of the group. For a number of reasons it
has been impossible to attempt a "complete” analysis of Antillean
lycaenid zoogeography. Instead, I have selected certain aspects of the
problem for study and presentation here, and these may be found at
the conclusion of the paper.
I. Nesiostfymon, new genus
Type species: Theda celida Lucas 1857
(ssp. shoumatoffi Comstock & Huntington 1943)
Hind wing tailed. Male fore wing with a diffuse scent patch, its scales
peripherally intermixed with ground color scales. Frons with appressed scales
only, lacking any erect, bristle-like scales. Eyes with sparse, short hair. Male
genitalia (fig. 3) : the whole genital ring dorso-ventrally attenuate, due ap-
parently to the ring being inclined (dorsal part to the rear) more than
usual, relative to the body axis; fakes of normal size, the fore arm straight
with a slight, short, gradual apical constriction; an abrupt angle at the elbow;
vinculum slender, sides subparallel for much of its length; saccus long, taper-
ing, slender, with a rounded, slightly inflated tip; valvae short (considerably
shorter, even, than the fore arm of the falx), completely fused with only a
slight apical notch to indicate the division, broadest half-way from base. Penis
long and slender, as usual, apically upcurving, with a single, simple exserted,
acuminate cornutus, lightly sclerotized; no ventral terminal keel or ventral (or
other exterior) teeth.
Remarks. Clearly related to Ministrymon Clench {leda Edwards, dytie
Edwards) and to Tmolus Hiibner (echion Linnaeus, azia Hewitson),
although well differentiated from both. It may be distinguished from these
two by the lack of bristly scaling on the frons, the more inclined genital ring
and the consequent attenuation of this ring dorso-ventrally, and in the very
short, completely fused valvae. With both it shares the single, simple acuminate
cornutus, lack of ventral keel on penis tip, the very long saccus and the rather
angular form of the fakes.
The brilliant blue above is more like Tmolus {echion) than Ministrymon,
as is the slightly more rounded wing shape and the diffuse scent patch.
The mainland (Mexico) species ter a Hewitson is quite possibly con-
generic but has not yet been examined critically.
252
CLENCH
/. Res. LepiJ.
II. Callophrys (Cyanophrys) crethona Hewitson
The genitalia of this species (fig. 1) indicate quite conclusively that
it is a true Cyanophrys. Like the pattern, however, they suggest that crethona
is more different from continental members of the subgenus than most of
the latter are from each other, thus implying a relatively great age to the
separation of crethona. Comstock & Huntington (1943: 123) state, however,
that it is closely related to an undescribed species from Mt. Roraima, Brasil,
a species I have not seen.
III. Pseudolycaena Wallengren 1859
Type species: Papilio marsyas Linnaeus 1758
Hind wing tailed, and with a tornal cleft. No scent patches. Frons with
appressed scales only, lacking any erect bristle-like scaling. Eyes with sparse,
short hair. Male genitalia ( fig. 2 ) : large, lateral hair pencils present, and
a dorsal chitinous piece of undetermined shape attached to anterior border
of tegumen; vinculum with posterior "shoulder” projection, vinculum strut
prominent, strongly curved into this projection; fakes rather strongly and
evenly curved; saccus eight to ten times as long as width at middle, the sides
subparallel, apex short, tapered, blunt; valvae rather short, broad, with lateral
denticulate wings, pointed apex, slight dorso-lateral tooth, mesial borders
divergent from before the middle. Penis with tip flared, definitely upturned,
with two terminal cornuti, the longer one with small, obscure, apical teeth;
terminal part of penis shaft with several short, external, longitudinal rows of
small teeth.
Remarks. Structurally the genus is allied to Atlides Hubner {halesus
Cramer) and Areas Hubner {imperialis Cramer), particularly the former.
All three of these genera share such significant characters as the extremely
long, slender saccus, compact valvae, apically flared and upturned penis, special
structural attachments on the anterior vinculum associated with the lateral
hair pencils; and all three have the curious and interesting tornal cleft on
the hind wing, as first pointed out by Godman & Salvin (1887: 15).
So far as I know the genus is monotypic, the sole species, marsyas Linn.,
with a Central American subspecies {damo Druce), a South American (nom-
inate marsyas) and the still little known Lesser Antillean subspecies, cybele
Godm. & Salv.
Fig. 1. Callophrys {Cyanophrys) crethona Hewitson, male genitalia
(slide C-705). Penis on left. This and the succeeding genitalia drawings
were made from mounts prepared as follows: first the penis was removed;
then the vinculum was cut through on the right side and the structures opened
out so that the inner sufaces are uppermost.' Drawings were made to show
the complete structures of the left side only (though usually both valvae are
drawn). Uncus, tegumen and falx are to the top right; saccus and valvae
to the left. This somewhat unorthodox procedure permits showing the struc-
tures with a minimum amount of distortion.
Fig. 2. Pseudolycaena m. marsyas Linnaeus, male genitalia (slide C-822).
Penis below, the central part of the shaft omitted, but with a scale to indicate
true relative length.
Fig. 3. Nesiostrymon celida shoumatoffi Comstock & Huntington, male
genitalia (slide C-7 08) . Penis on left.
Fig. 4. Heterosmaitia oppia Godman & Salvin, male genitalia (slide
C-823) . Penis below.
Fig. 5. Allosmaitia coelebs Herrich- Schaffer, male genitalia (slide
C-829) . Penis on left.
Fig. 6. Heterosmaitia bourkei Kaye, male genitalia (slide C-709). Penis
below.
Fig, 7. Heterosmaitia brescia Hewitson (?), male genitalia (slide C-826) .
Penis below.
2(4} .247-270, 196}
WEST INDIAN LYCAENIDAE
253
254
CLENCH
/. Res. Lepid.
IV. Heterosmaitia, new genus
Type species: Theda hourkei Kaye 1924 (Jamaica)
Hind wing tailed; males with androconial apparatus variable (see below).
Frons bristled to variable degree, Male genitalia (Figs. 4, 6, 7) : fakes with
tips prominently and abruptly constricted; vinculum with anterior triangular
projection (associated with the paired lateral eversible scale tufts or coremata);
saccus short, twice or less than twice as long as width at middle; valvae of
normal length, simple, mesial edges smooth; penis without ventral keel or
ventral teeth, more or less upturned apically, with two terminal cornuti. Males
all with a tornal red spot on hind wing above.
Remarks. The genus is mostly developed on the mainland, principally
in Central America. Investigation of potential relatives or possible members
is still incomplete, but studied members may be grouped as follows:
1. oppia group
Eyes with dense, long hairs. Frons with dense, erect, long bristles in
the median dark area; male fore wing without scent pad; hind wing with a
dark glandular patch at origin of Rs, with a few long, erea, specialized scales
from it near its posterior border; associated with this is a flat-lying, heavy
fringe of dark scales arising on the inner margin of the fore wing below,
direaed anteriorly; around this fringe the wing scaling is modified to form
a large opalescent-iridescent patch in Cu2-2A below cell, with a patch of dark
ochraceous mealy scales in the center. Male genitalia (fig. 4) : anterior
triangular process of vinculum small, close to saccus; valvae ventrally contiguous
to near their tips; penis strongly upturned apically; cornuti subequal in thick-
ness, with few or no apical teeth.
Includes only the Central American oppia Godman & Salvin.
2, hourkei group
Eyes with short, sparse hairs. Frons with bristles sparse, slight; male
with scent pad on fore wing at cell-end; no specialized scaling on fore wing
below or hind wing above. Male genitalia: anterior triangular process of
vinculum large, farther from saccus (more or less mid-lateral); mesial edges of
valvae divergent from before the middle; penis slightly upturned apically; one
cornutus apically much thicker than the other with many apical teeth.
Subgroup A. Saccus much less than half as long as valva; larger cornutus
with apical expanded part short, the expansion abrupt; penis slightly
upturned (fig. 6). Subterminal line of fore wing below rather well developed;
fuscous border of male fore wing above covers about half the cell-end-to-apex
distance.
Includes hourkei Kaye (Jamaica) and at least two Central American
species, one of which has been tentatively identified as guadala Schaus.
Subgroup B. Saccus much more than half as long as valva; large cornutus
with apical expansion gradual and rather l6ng; penis not at all upturned
(fig. 7). Subterminal line of fore wing below very faint; fuscous border of
male fore wing above covers about 1/3 the cell-end-to-apex distance.
Includes several apparently distinct species ranging from Mexico to South
America, including one tentatively identified as hrescia Hewitson. No West
Indian species are known.
V. Allosmaitia, new genus
Type species: Theda coelebs Herrich-Schaffer 1862 (Cuba)
Hind wing tailed. Eyes with rather long, dense hairs. Frons sparsely
bristled. Male without scent pad on fore wing but on hind wing with a
tuft of long, dark hairs arising from a small modified patch in the angle
between Sc and the costal border of the cell; associated with this there is on
the fore wing below a small, shining patch of modified scales on the inner
marginal area, near which vein 2A is slightly enlarged and scaleless, Male
2(4) ‘.247-270, 196)
WEST INDIAN LYCAENIDAE
255
genitalia (fig. 5): no triangular process on anterior border of vinculum;
fakes with gradually tapering tips; valvae exceeding long^ and slender, the
result of a greatly attentuated and produced terminal region, the mesial border
serrate terminally; penis apically very slightly and gradually upturned, with
two cornuti, one of which is distaliy enlarged, spatulate, and provided with
a flat terminal edge armed with a row of low teeth; saccus moderate in length.
Male above with no red spot at tornus. Fore wing of male with fuscous
border extending basad to cell-end.
Remarks. Clearly allied to the preceding genus, Heterosmaitia. differing
most notably in the long tuft of hairs on the hind wing aodroconial patch,
the absence of an anterior process on the vinculum, the extraordinarily pro-
duced valvae, the spatulate, truncate cornutus, as well as in the absence of a
tornai red spot on the male hind wing above.
Includes the following species: pion Godman & Saivin (Central America),
coelebs Herrich-Schaffer (Cuba), fidena Hewitson (Hispaniola, Puerto Rico).
In addition, the following species probably also belong but no specimens have
been available for study: piplea Godman & Saivin {=.subobscura Lathy 1)
(St. Vincent, Grenada, Dominica), panchaea Hewitson {=.scoteia Hewitson ? )
(Amazonas, Minas Geraes).
VL Eiectrostrymon Clench 1961
The West Indian species (save dominicana Lathy, which I have not
seen) are, according to both pattern and genitalia, more closely interrelated
than any is to endymion of the mainland, the type species and the only main-
land species I have so far investigated.
VII. ’Leptotes Scudder 1876
This genus is holotropical in distribution, being known in the Old
World chiefly under the name Syntarums Butler 1900, The American species
divide neatly, according to male genital characters, into three groups and
additional, co-equal, groups occur in the Old World. None of these groups
would seem to warrant being called even a subgeous. The three American
groups are:
(1) The Cassius group. This includes only cassim (with all its sub-
species) and a still undescribed species from the Colombian Andes.
(2) The marina group. Comprises marina Reakirt, perkinsae Kaye and
perhaps parrhasioides Waliengren from the Galapagos Islands. This group is
also represented by a number of species in Africa, Madagascar and associated
islands.
(3) The callanga group. This is perhaps the most primitive group in
the genus, and includes callanga Dyar, from the Peruvian Andes, and most
surprisingly wehhianus Bruile of the Canary Islands.
VIII. Tribe PLEBEJINI
The classification here is essentially that of Nabokov (1945), with his
subfamily Plebejinae demoted to a tribe, his relevant genera demoted to sub-
genera under Hemiargus, except for Pseudochrysops, which is distinct enough
in every way to be accorded full generic status.
IX. Subgenus Cyclargus
Nabokov groups these five species into two "superspecies” : ( 1 ) com-
prising dominica and mnmon; (2) comprising thomasi, woodruffi; eremhis.
Interesting confirmation exists in the sympatry (more or less) of ammon
and thomasi in the Bahamas; and Nabokov suggests that ehemhis may event-
ually be found in Cuba, sympatric there with ammon.
256
CLENCH
/. Res. tepid.
Present-day mean July temperatures (°F) in the West Indies.
2(4} -.247-270, 1963
WEST INDIAN LYCAENIDAE
257
X. Brephidium Scudder 1870
Carpenter & Lewis (1943: 392)- described Brephidium isophthalma
thompsoni in comparison with pseudofea Morr., under the mistaken, but then
prevaieot, opinion that is was nominate isophthalma. It is not distinguishable
from other Antillean populations of isophthalma.
Brephidium barbouri Clench was described as a distioa species, but later
was synonymized by Gillham (1955). In view of Gillham’s attitude to-
wards subspecies, and of the subtle and minute species differences in the
male genitalia of Brephidium which he may not have been able to appreciate,
as well as of the differences given in the original description, his action must
be restudied. Rindge’s remarks (1955: 14) are interesting in this connection.
ZOOGEOGRAPHICAL CONSIDERATIONS
The general pattern of lycaenid distribution in the Antilles is essen-
tially similar to that shown by most other groups of animals (cf.
Darlington^ 1957: 510). In brief, they constitute a depauperate fauna
of distinctly Central American aspect; they can be entirely accounted
for under an assumption of waif dispersal across water gaps similar to
those occurring today; the most striking endemics of the region occur
in Jamaica and Hispaniola; and despite the frequent development of
endemic subspecies in the different islands or island groups the insular
region as a whole presents a rather uniform, cohesive, facies. In con-
trast to some animal groups, however, the lycaenids of both southern
Florida and the Lesser Antilles contain such a sizable and unmistakable
West Indian component that for purposes of discussion these areas
must be considered integral, if peripheral, parts of the Antillean region.
The following discussion is not intended to be a full analysis of
West Indian lycaenid zoogeography. I have simply singled out for
discussion a few points that seem especially clear or particularly inter-
esting with emphasis on processes and events in the hope and belief
that these are of the most interest and value to others.
1. Late Pleistocene climatic changes
If there is one event that stands out more than any other as the
most important architect of the present distribution pattern of lycaenids
in the West Indies, that event would have to be the last (Wisconsin)
glaciation of the Pleistocene. It is well known, though still too little
appreciated by some zoogeographers, that during the times of glacial
advance temperatures dropped considerably not just at the ice front
but the world over. The actual amount of the drop undoubtedly varied
to some extent from place to place but it was on the order of approx-
imately 9° Fahrenheit (Flint, 1947: 455). Antillean lycaenids demon-
strate such a temperature drop quite unmistakably, and even indicate
certain of its details. Since summer temperatures in the West Indies
( fig. 8 ) are very little zoned there is no direct evidence in the lycaenids
whether or not these temperatures were affected by Wisconsin cooling^
^Though a certain amount of indirect evidence suggests that they probably were.
258
CLENCH
/. Res.
Lepid .
Present-day mean January temperatures (°F) in the West
2{4) -.247-270, 196}
WEST INDIAN LYCAENIDAE
259
but there is defioire indication that winter temperatures were lowered.
The amount of winter (January) mean temperature reduction that
best fits the present distributional evidence is approximately 8° F.
Interestingly only a very slight modification of the present January
isotherm contours (fig. 9) is required: a uniform reduction would
almost suffice. The needed modification is a slight shift to the east
of the "warm zone" which now lies over western Hispaniola, and of
the cooler zone which occurs just to the east over northeastern His-
paniola and western Puerto Rico, so that the first would lie squarely
over Hispaniola and the second directly over Puerto Rico.
The evidence that leads to these conclusions consists of three parts:
( 1 ) evidence for the reduction and its amount; ( 2 ) evidence for the
slight eastward zonal shift; and ( 3 ) evidence for the time when this
happened.
The reduction and its amount. Let us begin with the Antillean
element in peninsular Florida, which is dear-cut, of obvious derivation
(Cuba or the Bahamas) and, most importantly, is "open ended” ■ — ■
the northern limits of the ranges are determined by the direct inter-
action of the organisms and the environment, rather than being imposed
by an island shore.
Florida’s Antillean lycaenids may be divided into three groups
according to their relationships to their nearest relatives in the islands:
la. Undifferentiated. Chlofostrymon mae sites, Strymon manialis,
Leptotes cassius theonus.
lb. Subsped fically distinct. Eumaeus atala, Strymon acts, S. columel-
la, Hemiargus thomasi, H. ceraunus.
lc. Specifically distinct. Only Brephidium pseudo fea (whose prob-
ably postglacial invasion of the northern Bahamas may be ig-
nored for present purposes).
On the basis of their tolerance to January mean temperatures in
Florida they also divide into three groups but diffeerntly assorted:
lla. Northern boundary coincides more or less math 65° isotherm:
E. atala, C. maesifeSj S. acis, S. manialis, S. columella.
llb. Northern boundary coincides more or less with 62° isotherm:
L. cassius, H. thomasi.
lie. Northern boundary farther north than 60° isotherm:
H. ceraunus, B. pseudofea.
The hypothesis which best explains the above facts is that at some
past time conditions in Florida were measurably cooler than now; that
during this cooler time and because of it the greater part of the An-
tillean element was absent; and that subsequently conditions warmed
to about their present state, permitting the island forms to invade
the peninsula and establish themselves, creating the present fauna.
Important in this connection is the degree of differentiation of these
260
CLENCH
/. Res. LepiJ.
invaders. This is discussed more fully below and all that need be said
here is that with the exception of pseudo fea (Ic above) they show
a fairly consistent degree of differentiation ("slight” subspecies to
undifferentiated) of a small order of magnitude. This argues for a
nearly simultaneous entry into Florida not very long ago (as opposed,
for example, to random arrival and establishment at intervals over a
long period of time. )
A drop of twenty degrees would be required to completely elim-
inate all the West Indian species now in Florida because of the con-
siderably more cool tolerant group lie above. Such a reduction is
patently absurd for it would also have eliminated virtually all the
lycaenids in the islands too. A much more realistic 8° would suffice
to lower the mean January temperature of warmest Florida from its
70° tp 62°, probably sufficient to wipe out all but the rebellious Ik
species and further south not too severe for survival on some islands.
Taking into consideration the distinct possibility that some species,
particularly those of group lib, may have become adapted in Florida
to somewhat cooler conditions than they could tolerate when they first
arrived (for several reasons more likely on the mainland than in the
islands), a more conservative 6° might be suggested. The higher
figure, however, is preferable because of what apparently happened
in the islands: a drop of only 6° would be insufficient to explain the
pattern of survival which is indicated there (discussed more fully be-
low). Nonetheless this reasoning suggests that in the absence of the
stronger selection pressures of the mainland the minimum tolerance
of Antillean lycaenids was about 66° mean January temperature.
The eastward zonal shift. A reduction of 8° in the present Antillean
January isotherm of 74° would bring it down to the suggested critical
value of 66°, and this isotherm may be imagined as roughly dividing
the insular region into two areas: a northern one which in Wisconsin
time had winter temperatures below the critical value; and a southern
one which had temperatures above this value and hence presumably
could have served as a refuge area during that glaciation.
An island which today harbors one or more endemic full species
can be presumed to have maintained at least those species through
Wisconsin times. Jamaica has five such endemics, Hispaniola three,
Cuba three (including E. atala which, though not endemic, must have
survived the glaciation there). Puerto Rico and the Bahamas have
none. There are also several significant endemics in the Lesser An-
tilles, but distributional knowledge is insufficient to discuss them in
detail. It is legitimate to conclude from this that Jamaica, Hispaniola,
and the Lesser Antilles must have had adequate winter temperatures
during Wisconsin time while Puerto Rico and the Bahamas did not.
Cuba, on this basis, probably had adequate conditions but only marg-
inally so.
2(4) -.247-270, I96>
WEST INDIAN LYCAENIDAE
261
The present January isotherm of 74° (hypothetical Wisconsin 66°)
comes close to dividing the Antilles in just that way. It passes barely
through southernmost Cuba, consistent with the marginal conditions
just suggested; Jamaica and the Lesser Antilles lie wholly below it
and Hispaniola nearly so, in harmony with the conditions suggested by
their relatively large number of endemics.
But as it stands it is not perfect. Marginal conditions, according to
the present configuration of the isotherm, should have prevailed in
the southern Bahamas and in Puerto Rico, whereas there is no evidence
that any lycaenids survived in either. Adjustment of the configura-
tion to conform with the endemics (and lack thereof) is, however,
minor: push it southward where it intersects the Bahamas'^ and nudge
it a little to the east where it crosses Hispaniola and back again over
Puerto Rico. The results are shown in fig. 10.
The time it happened. The best source of information on this
point is the degree of differentiation that has taken place between the
forms in Florida and their nearest insular relatives (see groups la, Ib,
above). The maximum differentiation is shown by the members of
Ib, all of which are "slight” subspecies of insular species. This degree
of divergence is typical of what is imputed to post-Wisconsin effects
in many groups of animals (not just lycaenids, or even just butterflies),
and in places where glacial influence is more directly ascribed to the
differentiation. Little more need be said: by their degree of divergence,
the concordance of several forms, and their presence in a place where
conditions were appropriate, they indicate arrival in Florida, either from
Cuba or the Bahamas, closely following the warming temperatures that
accompanied recession of the Wisconsin ice sheet far to the north.
As to the undifferentiated species in group la, there are two explana-
tions. They might be late arrivals and accordingly have had too little
time to become differentiated; or they might have arrived with the
others but because of a much lower evolution rate have simply failed
to diverge. I incline to the latter possibility for a number of reasons:
First, these same species show similar effects among the islands:
no divergence between islands where other species have become post-
glacially divergent, sometimes (e.g. C. maesites) showing subspecies
only where a separate glacial refuge can be invoked.
Second, when the temperatures were warming at the end of Wis-
consin time the northerly islands must have presented a sort of ecolog-
ical vacuum extremely receptive to immigrants from the south. Since
then, however, it is likely that the hypothesized "closed association”
or "closed community” effect has become operative. This hypothesis
little push only; a very few butterflies, Eurema chamberlaini Butler being
the most conspicuous, seem to have been able to persist in these islands dur-
ing the Wisconsin.
262
CLENCH
/. Res. Lepid.
Fig. 10. Assumed mean conditions for January during Wisconcin glacial
maximum. Line represents approximate position of 66 °F isotherm (position
and configuration based on present day January isotherm of 74° with modifica-
tions: see text). Shaded areas represent presumed refugia during Wisconsin
2(4) •.247-270, 196}
WEST INDIAN LYCAENIDAE
263
(cf. Weaver & Clements, 1938: 171) holds that in a climatically stable
situation communities so adequately fill the available niches . that they
tend to impede the establishment of immigrants from without. It is,
of course, not an absolute condition but a tendency. It must, however,
become increasingly effective with decreasing distance from the pros-
pective source of the immigrant. Between the West Indian islands,
for example, it seems probable that movement between adjacent islands
is sufficiently frequent that those forms capable of becoming establish-
ed have already done so long ago while those incapable continue to fail.
Third, statistically it is more logical to expect a variety of evolution
rates in a group than a single rate. There is no reason whatever to
expect a form to subspeciate simply because it has the chance. Florida’s
Antillean lycaenids present an entirely reasonable pattern of some low
rates, some higher rates (time has been insufficient to distinguish
clearly between "high” and "not so high” rates, however).
Resume. It would be well to recapitulate the events discussed
above, putting them in chronological order.
( 1 ) The advance of the Wisconsin ice sheet was accompanied
all over the world by a marked drop in temperature. In the West
Indies the effect was to reduce mean January temperatures by about
8°F., the effect, if any, on summer temperatures being indeterminate.
(2) The result of this was to extirpate completely the Antillean
lycaenids in Florida (except Brephidium pseudofea) , the Bahamas and
Puerto Rico, and most of them in Cuba. Survival was possible on a
large scale only in Jamaica, Hispaniola and in the Lesser Antilles (the
latter discussed below ) .
(3) When the Wisconsin sheet retreated, some 12,000 years ago,
conditions returned to approximately their present state and concurrent-
ly many species that had been confined to the three or four refugia
just mentioned spread out rapidly, occupying all islands (more or less)
capable of supporting them.
(4) Since that time certain of these species have evolved, or have
continued to evolve, producing subspecific differentiation among the
islands. Others with a slower evolution rate have remained visibly
undifferentiated.
It may be of interest to summarize the probable or possible ly-
caenid faunal composition of the four principal refugia at the height of
the Wisconsin glaciation. The evidence allocating a particular species
to one or another refugium is various. Endemic full species are, of
course, immediately assignable. But assignment of others is based on
the most likely refugium, granted the foregoing sequence of events.
Species that today show among the islands subspecies of about equal
divergence one from another (e.g., the subspecies of E. angelia) are
assumed to have spread out from a single refugium. Those such as
S. acis that show two groups of subspecies suggest double refugia with
subspeciation already in progress between them during Wisconsin time.
264
CLENCH
/. Res. Lepid.
1. Cuba. E. atala, A. coelebs and H. ammon are endemic there
or nearly so, and apparently survived the Wisconsin there. A total of
three species.
2. Jamacia. C. crethona, H. bourkei, E. pan^ L. perkinsae and H.
dominica are all endemic full species and must have survived the
Wisconsin in sUu\ in addition the following were likely: S. ads (an-
cestor of sspp. bartrami, casasi, gossei, armouri) , S, martialis, S. colum-
ella ( ancestor of sspp. modesta^ cybira ) , L. cassius theonus, H. ceraunus
(ancestor of sspp. filenus, ceraunus, antihubastus) , B. exilis isophthalma.
A total of 11 species listed.
3. Hispaniola. S. cristophei, S. toussainti, P. bornoi are all endemic
there today. Additionally the following were likely: A. fidena, S. acts
ancestor of sspp. acts, mars, petioni) S. columella (ancestor of sspp.
columella, arecibo ) , S. limenia, E. angelia, H. thomasi. A total of nine
species listed.
4. Lesser Antilles. A. piplea, S. antigua, E. angerona, E. dominica
are all endemic there today. The following were likely in addition:
P. marsyas cybele, S. buhastus ponce, L. cassius catilina, L. cassius chad-
wicki, H. hanno watsoni, H. woodruffi. A total of nine species listed,
one of them in two subspecies.
The following species were undoubtedly present on one or more of
these refugia but the particular one (or ones) cannot now be deducted
from available . data : N, celida; S. bazochii; C. ma^ sites (ssp. clenchi
probably was present in the Lesser Antilles but the refugium of m.
maesites cannot be determined); C, simaethis (ssp. jago w’as probably
present in the Jamaica refugium but the refugium of s. simaethis is not
determinable.
In concluding this subject it may be of interest to add some observa-
tions on three related matters.
First, the Wisconsin extirpation of the Antillean lycaenids in Florida,
the Bahamas, Puerto Rico and at least partially in Cuba, does not imply
that these areas were without Lycaenidae during the glaciation. Florida,
to judge by availability and systematic relations today, must have had a
sizeable number of species; but which of these, if any, might have
invaded the islands themselves is not known, nor is it ever likely to be
since they have now all disappeared. From this last I should guess that
the putative transient boreal element could not have been particularly
large in the islands, or some would surely have managed somehow to
survive into the present. A few species in other butterfly families,
however, suggest just such a history. Phyciodes phaon Edv/ards (Nym-
phalidae) is a possible example.
Second, although it cannot be invoked in explanation of any known
lycaenid distribution, the matter of altitudinal range deserves brief
notice since it is definitely involved in the distribution of certain other
butterflies. A species adapted to conditions found above an elevation
of roughly 2000-2500 feet or so could have survived the suggested
2(4)t247.270. 196}
WEST INDIAN LYCAENIDAE
265
glacial temperature drop more or less in place simply by moving down-
ward. Such groups of species as Calisto (Satyridae), Atlantea (Nym-
phalidae ) (Pieridae) and Hymenitis (Ithomiidae) prob-
ably owe their preservation through the Wisconsin glaciation to this
factor.
Third, there are several groups of related species in the islands
whose affinities suggest that they arose within the island area by a
process analogous to that which has produced the now observable sub-
species in many species. The Antillean species of Electfostrymon form
one such group; the subgenus Cydargus another; the Antillean Alios-
mdtia probably yet another. Each of these groups suggests dispersal
through the islands from a refugium (or original beachhead of estab-
lishment in the islands) closely following a glacial recession. The
degree of divergence in these instances and the subsequent history of
several of the component species imply forcibly that a glaciation older
than the Wisconsin was responsible, perhaps the next earlier (Illinoian).
2. Mainland colonization from the islands
Under this heading I propose to unite the discussions of three differ-
ent examples, all of which share the possibility that they have colonized
the Central American mainland from the islands. This seems to be a
novel idea: at least I know of no prior suggestion of it in any group.
It is hard to tell though whether this is due to the probable rarity of
the phenomenon or to such preoccupation with the many problems
associated with invasion from the mainland that the idea of a reverse
colonization simply never occurred to anyone. In any case there are
three instances in the lycaenids where this seems to be indicated. Par-
enthetically it may be noted that of roughly 20 successful migrations
between islands and mainland (other than Florida) needed to account
for the Antillean Lycaenidae, three (15%) are of this abinsular type,
the remaining 17 being more conventional adinsular colonizations. If
the lycaenids represent a fair zoogeographic sampling of the island
fauna, 15% is quite large enough to make abinsular colonizations worth
looking for in other groups.
1. St-rymon columella. This species does not occur in South Amer-
ica but it is present in Central America (including Baja California and
some of the offshore Pacific islands) where it ranges north into south-
ern United States, and it occurs on all the major islands and island
groups of the Greater Antilles, There are several related species in
the West Indies {antigua, toussainti, cristophei, limenia) , while in
South America occur still other allied species. This pattern suggests
that there was an early colonization of the Antilles, probably from
South America, followed by speciation within the islands that resulted
eventually in the formation of columella. Then, most likely during
the Wisconsin, the latter crossed over into Central America. With
266
CLENCH
/. Res. Lepid.
the recession of the Wisconsin sheet, in addition to spreading out from
its island refugia it also spread, in Central America, to Baja California
and to some of the islands off the West Coast.
The remaining two examples are even more unusual in that they
show definite evidence of having come originally from Africa. So far
I am aware these two, the Leptotes marina group and Brephidmm, are
the only New World Lycaenidae to which an African origin can be
attributed. It has been suggested that Zizula cyna Edw. (purely con-
tinental from southern United States south to Brazil and Bolivia) is
such an African derivative but fuller investigation does not support
the contention. Z, cyna is a distinct New World species; its closest
ally, the polytypic Z. gaika Trimen, ranges from Africa to Australia.
Comparison of cyna with various gaika populations show a distinctly
greater resemblance to those from eastern Asiatic localities than to
those of Africa. The only conclusion possible on this evidence is that
cyna was derived from Asia via the Bering Strait route and not from
Africa.
2. Leptotes marina group. As pointed out above (note no. 7) this
group consists of a number of species in Africa, Madagascar and several
of the islands in the western Indian Ocean as well as the three New
World species, perkinsae ( Jamaica ) , marina ( Central America ) and prob-
ably parrhasioides Wallengren (Galapagos Islands).^ The marina group
occurs nowhere else but related groups also occur in Africa. The pres-
ence of members of the group on so many remote oceanic islands is
dramatic evidence of a considerable propensity for colonizing across
large water distances so it would be by no means impossible for a
hypothetical ''marina group ancestor” to have made the long voyage
from Africa to the New World. The real question is the site of its
landfall. On the face of it South America would appear to be the best
answer by far; it is nearest to Africa and it is the largest available
target for a random trajectory to hit. The evidence against this possi-
bility, however, is to me persuasive though admittedly tenuous.
First, the present distribution of the New World members of the
group is distinctly northern tropical, almost subtropical. Only parr-
hasioides is truly equatorial (in latitude but not in climate).
Second, for some unknown reason the tropics generally are not very
favorable for the Plebejinae. This is least true in the East Indies, most
true in the New World. It is therefore distinctly possible that a form
of this subfamily arriving in eastern South America would be poorly
equipped a priori to compete and establish itself there. In the West
^Comstock & Huntington (1943:95) cite marina from Chile and the Galapagos.
It is hard to question the accuracy of such meticulous observers but it is even
harder to accept these records. Either mislabelling or misdetermination would
seem to be involved here.
2(4)1247-270, 1963
WEST INDIAN LYCAENIDAE
267
Indies, on the other hand, conditions appear to be much more favorable
to the subfamily. Leptotes, Hemiargm and Pseudochrysops flourish
there and together make up a larger percentage of the lycaenid fauna
than in any other tropical area in the Americas.
Third, comparison with Brephidium (which presents at- least one
additional type of evidence) shows a strongly similar pattern in many
respects, and each thereby lends support to the other.
The tentative conclusion is that the ancester of the New World
members of the marina group crossed from Africa to the West Indian
region where it became established; that from there it successfully
invaded the Central American mainland; and then spread from there
to the Galapagos. It is impossible, at least with the information avail-
able now, to guess when this might have happened. Diversification
of the group in the New World has left no tangible clues to evolu-
tion rates. One might say that if evolution rates in the group were
more or less typical, then the Atlantic crossing might have been made
in mid-Pleistocene or thereabouts but that is completely without doc-
umentation,
3. Brephidium. There are many points of similarity to the marina
group in the distribution pattern of this genus, and just enough differ-
ences that, if they share a similar history, one tends to complement the
other in evidence. There are three known species of Brephidium.
(omitting B. barhouri because of insufficient information) :
B. pseudofea Morr., confined to Florida and Georgia with a single
recently discovered colony (Rindge, 1952: 13) in the northern Bahamas.
B. exilis Bdv., widespread in the New World with these known
and very distinct subspecies: nominate exilis from Oregon and Nebraska
(probably not overwintering) south so far as now known to eastern
Texas and on the Pacific Coast to southern Baja California and to
Mazatlan (Sinaloa, Mexico) at least; a still undescribed subspecies
from coastal Yucatan; probably one or more other subspecies down to
northern Venezuela, the limit of the known continental distribution
of the species; e. isophthalma from the West Indies.
B. metophis Wallengren in South Africa, chiefly in the western
Cape Province and western Orange Free State, more or less sympatric
there with the related monotypic genus Oraidium {harberae Trimen).
See Swanepoel (1953: 80-81) for information on these two species.
No other members of the genus and no other closely related genera
are known anywhere.
The northern range of exilis on the mainland and its nearly com-
plete absence from South America implies a northerly source, further
supported by the presence of ssp. isophthalma in the Greater Antilles.
The distinctness and the presence in Florida of pseudofea constitute
strong presumptive evidence that it is there a survivor of an earlier
invasion from the islands, again implying a longer occupancy by the
genus in the Antilles than on the mainland, where only one species is
known.
268
CLENCH
/. Kes. Lepid.
These points lead to the conclusion that an ancestral Brephidium
crossed from Africa to the West Indies and became established there;
then later spread to Florida to form pseudofea; and still later crossed
to the mainland to give rise to the several subspecies of exilis now
known there.
We are in a somewhat better position in Brephidium than in the
marina group to date these events, however roughly. The presence of
isophthalma on several Greater Antillean islands without apparent sub-
speciation is evidence that the evolution rate oiexilis is rather low,
requiring more than post- Wisconsin time to produce visible sub-
speciation. We are, therefore, led to conclude that the invasion of the
mainland, where several distinct subspecies now occur (all different
from the insular subspecies as well), preceded the Wisconsin; and that
the origin of Floridian pseudofea must be much older still. The whole
New World history of the genus could have been written since the
onset of the Pleistocene, and may well have been, but it could easily
have been even older.
In passing it may be noted that this suggestion of an African
origin of New World Brephidium carries with it the likelihood that
at the time of the Atlantic crossing Brephidium in Africa enjoyed a
wider, or at least considerably different, range than it does now.
Possibly it was then an inhabitant of the savanna regions of the interior
of Upper Guinea.
3. The Lesser Antilles fauna
Our knowledge of the lycaenids of this curious chain of islands
appears to be still fragmentary, but there are two points at least which
warrant brief remarks and for which the distributional evidence seems
adequate.
Mention has already been made of the likelihood that the Lesser
Antilles served as a Wisconsin refugium. This refugium, however, ap-
pears to have functioned much less as a source of postglacial spreading
than either of the other two primary refugia, Jamaica and Hispaniola.
These facts are relevant: (1) the Virgin Islands are faunistically a part
of the Lesser Antilles (cf. Comstock & Huntington, 1943: 121); (2)
of the 12 lycaenid species regularly occurring in Puerto Rico, the first
"port of call" for a potentially expanding Lesser Antillean refuge fauna,
three (C, maesites, C. simaethis, N. celida) must be omitted for un-
certainty of origin; three {S. huhastus, H. hanno, H. woodruff i) are
clearly derived from the Lesser Antilles in postglacial times while the
remaining nine are apparently derived from Hispaniola; ( 3 ) St. Kitts
shares most, perhaps all, of its fauna with the Virgin Islands, but the
remaining Lesser Antilles do not do so to the same extent. These three
points suggest that the failure to expand postglacially may be more
apparent than real. The problem would be neatly solved by making
2(4) -.247-270. 196}
WEST INDIAN LYCAENIDAE
269
the assumption that the Virgin Islands shared Puerto Rico’s low temper-
atures during Wisconsin times and were therefore not inhabited; that
when recession of the Wisconsin sheet began, invasion of Puerto Rico
started from Hispaniola while simultaneously the Virgin Islands were
being invaded from St. Kitts. This would explain not only the predom-
inantly Hispaniolan character of the Puerto Rican fauna and the Lesser
Antillean character of the Virgin Islands fauna but also the apparent
failure of the Lesser Antillean refuge fauna to disperse; for under
this hypothesis it would not have lacked capacity to disperse but simply
did not have the opportunity to spread so far because of the opposing
expansion of the Hispaniolan refuge elements.
Strikingly apparent is another aspect Of this Lesser Antillean fauna:
its integral relation to, and derivation from, the fauna of the Greater
Antilles. Fourteen species are known from the islands, from the
Virgin Islands to Grenada inclusive. Of this number one remains an
enigma as to its origin (S. mfofusca); one is indeterminate and could
have reached the Lesser Antilles from either the north or the south
( C. simaethis ) ; three are of unquestionable South American origin
(P. marsyas, S. huhastus, H. hanno) , and L. c. cassius may also have
invaded the islands from South America as far as Martininque (only
a single male is known from the island ) . The remaining nine species
are of obvious Antillean derivation: A piplea, S. acts, S. antigua, E.
angelia, E. angerona, E. dominicana (probably), C. maesites, H. tvood-
ruffi^ L. cassius (sspp. catalina and chadwicki) . Counting only those
of reasonably assured origin shows that the Lesser Antillean fauna is
about 75% Greater Antillean in origin, 25% South American.
4. The age of the Antillean lycaenid fauna
Throughout the preceding discussion attempt has been made
wherever possible to apply some sort of age estimate to the events set
forth. By extending analogous reasoning as well to species not specifi-
cally discussed we reach an interesting conclusion 'Concerning the age
of the lycaenid fauna now in the Antilles. At least two glaciations
are required for certain species groups {Brephidium, Electrostrymon,
Cyclargus); no member of the fauna is so remote systematically from
its nearest continental ally that the differentiation could not have
occurred since the onset of the Pleistocene. One may conclude from
this that early Pleistocene, more or less, is the most likely time for
the initiation of events that culminated in today’s fauna. If evolution
rates were substantially less than is here estimated, then of course this
time would need to be pushed back accordingly.
In brief there is no firm evidence of the time this fauna first began
to invade the islands, but contributing data in the form of systematic
affinities and suggested or implied evolution rates in "typical” lines
lead to the conclusion that early Pleistocene is the most likely time.
270
CLENCH
/. Res. Lepid.
BIBLIOGRAPHY
BATES, M., 1935. The butterflies of Cuba. Bull. Mus. Comp. Zool. 78:
63-258, 23 figs.
CARPENTER, G. D. H., and C. B. LEWIS, 1943. A collection of Lepidoptera
(Rhopalocera) from the Cayman Islands. Ann. Carnegie Mus. 29: 371-
396, 9 figs.
CLENCH, H. K., 1961. Theclini (pp. 177-219) in P. R. and A. H. Ehrlich,
How to know the butterflies. Dubuque, Iowa (W, C. Brown), [8] -f-
262 pp., 525 figs.
COMSTOCK, W. P. and E. I. HUNTINGTON, 1943. Lycaenidae of the
Antilles (Lepidoptera, Rhopalocera). Ann. New York Acad. Sci. 45: 49-
130, 2 text figs., 1 plate.
DARLINGTON, P. J., 1957. Zoogeography: the geographic distribution of
animals. New York (Wiley), xi, 675 pp., 80 figs.
FLINT, R. F., 1947. Glacial geology and the Pleistocene epoch. New York
(Wiley), ix, 589 pp., 88 text figs., 6 plates.
GILLHAM, N. W., 1955. Brephidium barbouri Clench a synonym of Bre-
phidium exilis isophthalma (Herrich-Schaffer) (Lepidoptera: Lycaenidae).
Psyche 62: 34.
GODMAN, F. D., and O. SALVIN, 1879-1901. Insecta, Lepidoptera-Rhopal-
ocera, in Biologia Centrali- Americana. Vol. 1 (1879-1886, 1901). xlvi,
487 pp.; vol. 2 (1887-1901), 782 pp.; vol. 3, 112 plates.
NABOKOV, V., 1945. Notes on neotropical Plebejinae (Lycaenidae, Lepid-
optera). Psyche 52: 1-61, 1 text fig., 8 plates.
RINDGE, F. H., 1952. The butterflies of the Bahama Islands, British West
Indies (Lepidoptera). Am. Mus. Novit. #1563, 18 pp.
— — 1955. The butterflies of the Van Voast- American Museum
of Natural History Expedition to the Bahama Islands, British West Indies.
Am. Mus. Novit. #1715, 20 pp., 8 figs.
SWANEPOEL, D. A,, 1953. Butterflies of South Africa. Cape Town (M.
Miller), 320 pp., 8 halftone, 17 colored plates.
WEAVER, J. E., and F. E. CLEMENTS, 1938. Plant Ecology (2nd Edition).
New York (McGraw-Hill), xxii, 601 pp., 271 text figs., frontispece.
Journal of Research on the Lepidoptera
2 (4) :271-280, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
(cj Copyright 1964
THE SYNONYMY, VARIABILITY
AND BIOLOGY OF LYCAENA NIVALIS
E. J. NEWCOMER
1509 Summitview, Yakima, Washington
Lycaena nivalis Bdv. is a very interesting species in several
ways. (1) Its synonymy has been somewhat confused; (2) it is
quite variable, which may explain the confusion of names; and
(3) its life history was described more than 50 years ago in the
Canadian Entomologist, but this fact has gone unrecognized,
again because of confusion of names.
L. nivalis^ and its subspecies browni Dos Passes, occurs,
according to Clench (1961), "from southern British Columbia
south to Colorado, Utah and the middle Sierras of California.”
Llewellyn Jones (1951) says it is found in British Columbia in
the Southern Interior and the Kootenays, in other words, the
entire area between the Fraser River and Alberta north to about
latitude 52°. In Washington the writer has taken it in Okanogan,
Yakima and Klickitat Counties, and it is particularly abundant
in Yakima County. Leighton (1946) records it from Leaven-
worth, which is in Chelan County, and it should also occur in the
intervening Kittitas County and perhaps elsewhere in the state.
In Oregon the writer has taken it on Mt. Ashland, Jackson
County; at Tombstone and Lost Prairies, Linn County, which are
just west of the Cascade summit at Santiam Pass; in the Ochoco
and Maury Mountains, Crook County; and in the Warner Moun-
tains, Lake County. In California he collected it years ago just
north of Lake Tahoe and more recently just west of Ebbet Pass,
Stanislaus County, and in the Warner Mountains, Modoc County.
Comstock (1927) says it occurs "at high elevations throughout
the Sierras.” Brown (1957) records the species from northern
and western Wyoming and northern Utah, and the subspecies
browni from southern Idaho, northwestern Colorado and south-
eastern Wyoming. The writer has taken it at Aspen Grove,
Utah County, Utah.
271
272
NEWCOMER
/. Res. Lepid.
Fig. 1. Lycaena from Oregon and Washington.
a. Typical Lycaena nivalis. Maury Mts., Crook Co., Oregon, b. Similar but
females melanic, and black spots below more pronounced. Camas Creek,
Warner Mts., Oregon, c. Typical L. n. hrowni. Bear Canyon, Yakima Co.,
Wash. d. Similar below, but colors more intense; females with very little
melanism. Cascade Mts., Oregon.
2 (4):271-280, 1963
LYCAENA NIVALIS
273
SYNONYMY
Edwards ( 1877 ) listed nivalis as a synonym of mariposa and
ianthe as a synonym of zeroe, and this synonymy was followed
by Dyar (1902). Barnes & McDunnough (1917) listed zeroe as
a synonym of mariposa and ianthe as a synonym of nivalis, and
this synonymy has been used by Comstock ( 1927 ) and Holland
( 1931 ) . Holland’s original Butterfly Book ( 1904 ) did not recog-
nize nivalis, ianthe or zeroe at all. Wright (1907) figured what
is certainly nivalis but called it zeroe.
The writer has referred to the original descriptions of ianthe,
nivalis and zeroe\ and it is difficult to see how some of this
synonymy came about. Edwards (1871), in describing ianthe,
says in part: “Underside pale buff much powdered at base with
grey; primaries have a marginal series of rounded brown spots,
obsolete apically, suffused near inner angle with fulvous; a
median tortuous row of brown spots or points; a bar on arc,
a large round spot below cell. Secondaries have a marginal
series of orange crenations, obsolete toward outer angle; a
medium row of black dots; an obsolete discal streak; a point in
cell and another below.” This certainly describes nivalis, so
that Barnes & McDunnough were correct in making ianthe
synonymous with that species.
Boisduval’s (1869) description of nivalis is in Latin and
French, and it may be translated into English about as follows:
Upper side of wings darkly fulvous, middle lunules black, shot with
iridescent purple; posterior marginal band fulvous; fore wings below pale
ochraceous with black spots; hind wings pale, lunules ferruginous.
It looks something like our Mere but is very small. The fore wings of
the male are a smoky fawn with a beautiful violet reflection. Each wing
is marked in the middle with a blackish lunule and with several obsolescent
spots faintly visible; below there is a small crenelated band at the margin of
ferruginous fawn, bordered behind by black lunules. Below, the fore
wings are yellow-ochre, very pale, with black spots as in our species; the
secondaries are rosy-yellow, a little glaucous at the base, with some small
black spots scarcely indicated, and a series of marginal, obsolete fawn-
colored lunules. The female is a clear yellow-fawn, spotted as in other
species of the group. Below, it does not differ from the male. This species
is very distinct, and similar to helloides. It is found in the Sierras.
Nivalis, meaning “belonging to the snow,” is an appropriate
name for the species, as it occurs only in areas where there is
considerable, and sometimes very heavy, snowfall.
Boisduval’s (1869) description of zeroe, again in Latin and
French, may be Anglicized as follows:
^The latter two courtesy of F. Martin Brown.
274
NEWCOMER
/. Res. Lepid.
Wings darkly fuscous, with purple iridescence, middle unules margin-
ally dark fuscous; fore wings with black spots. Fore wings below ochraceous
with black spots; hind wings cinereous with thin black spots.
Something like our hiere but smaller and in size similar to the pre-
ceding. The fore wings of the male are of a brownish tinge washed with
fawn and with a violet reflection, and a small discoidal, blackish lunule at
the edge; the wings above are marked with black spots; below in the anal
angle is a small area with a trace of fawn lunules. The hind wings above
are pale yellow-ochre, with black spots; below ashy gray with black spots
in the lunules, transversely arranged; on the hind edge there is a series of
sagittate whitish spots cut with black. The female is a lively yellowish
fawn, spotted with black as in other species. This pretty species inhabits
the high mountains of the Utah border in June and July.
The description of zeroe follows that of nivalis in Boisduval’s
paper, so that his statement that zeroe “se rapproche par la taille
de I’espece precedente” is comparing the size of the two. And it
is obvious that this description of zeroe would fit mariposa quite
well, as the secondaries of the latter are certainly “ashy gray”
below. One cannot help wondering if imperfect translation of
these descriptions caused Edwards to list nivalis as a synonym
of mariposa, and Wright to figure nivalis and call it zeroe.
In view of the above information the present synonymy
appears to be correct.
VARIABILITY
The genus Lycaena includes a number of species that are
quite variable as attested by the names given to subspecies as
well as the names that have become synonyms. L. nivalis is no
exception and its variability may well be due to its occurrence
under widely differing climatic conditions. This is made pos-
sible by the distribution of its major food plant. Polygonum
douglasii, in both the Transition and Boreal Zones, at elevations
of at least between 2,000 and 7,500 feet, and where annual pre-
cipitation ranges from about 15 inches to 60 inches or more.
Brown (1957) reports that in Colorado nivalis frequents
sage-brush flats and is rarely found along streams. In Oregon
the writer has taken it at 7,000 feet on the slopes of Mt. Ashland;
in a lush meadow at 4,000 feet in Linn County; in rather dry
yellow-pine areas at 5,000 to 6,000 feet in the Ochoco and Maury
Mountains in Crook County; and in somewhat similar areas at
6,000 to 7,500 feet in the Warner Mountains in Lake County.
In Washington it may be found in canyons at relatively low
elevations and also on mountain slopes and in high meadows.
So variability is probably inevitable.
2 (4);271-280, 1963
LYCAENA NIVALIS
275
Material from Tombstone and Lost Prairies has a very in-
tense coloration, the yellow and pink of the under side being
brilliant, and the fawn color of the upper side of the females in
very sharp contrast to the fuscous spots and margins. The same
coloration occurs on material from Mt. Ashland, where pre-
cipitation would.be at least 40 inches, and to a considerable ex-
tent on material from Harts Pass in northern Washington where
the precipitation is high.
Specimens from the Ochoco and Maury Mountains, on the
other hand, have a much less intense coloration, the ground color
beneath, even in fresh specimens, being a washed-out light
cream, with a tinge of pinkish on the secondaries, but no evi-
dence of yellow. The color of the males above is somewhat less
intense, and the females have a considerable smoky suffusion
above. Here the spring rainfall is light and the annual not over
20 inches. Material from the Warner Mountains is similar to
that from the Ochocos, except that the females are more melanic.
Precipitation there is probably 20 to 25 inches.
Specimens from Satus Pass and Bear Canyon have rather
contrasting yellow and pink coloration below, not as intense as
on specimens from the Oregon Cascades, but much brighter
than on the Ochoco material. Satus Pass females have perhaps
less smoky suffusion, on the whole, than those from Bear Canyon.
It should be pointed out that in the areas where intense color-
ation occurs, there would be less sunlight than in the areas
where the colors beneath are less intense, and this may be a
factor in determining the intensity of the colors.
Mean temperatures have been examined, particularly those
of May and June, which might be expected to have the most
effect. The highest temperatures occur in the southern part of
the area and they decrease to the north. This does not match up
with the difference in the coloration. And, as can be seen in
the table, elevation is probably not a factor.
Lacking experimental data, no explanation is evident for the
color variations found in nivalis, other than those suggested here.
SUMMARY OF VARIETIES
There are four rather distinct varieties of nivalis in the area
under discussion:
I. Typical nivalis, mostly pinkish below with some black
spots on the secondaries; the females with much yellow or
276
NEWCOMER
/. Res. Lepid.
orange above. Sierra Nevadas, California, and Ochoco and
Maury Mountains, Oregon. Fig. la.
2. Similar, but females very melanic above and black spots
on secondaries below more pronounced. Warner Mountains,
Oregon. Fig. lb.
3. Typical browni^ females with some orange above to very
melanic; contrasting pink and yellow below and black spots on
secondaries much reduced or absent. Utah to Washington.
Fig. Ic.
4. Similar below, but colors much more intense, the pink
contrasting strongly with the yellow; black spots reduced or
obsolete; the females above with very little melanism, the
orange ground color contrasting greatly with the fuscous spots
and margins. Cascade Mountains of Oregon, specifically Linn
and Jackson Counties. Fig. Id.
The writer is not a taxonomist, but if browni is a valid sub-
species, it would seem that this Cascade Mountain variety could
legitimately be named as another subspecies, particularly as the
conditions under which it occurs are very different from those
elsewhere.
LIFE HISTORY
The writer described the early stages of nivalis more than
50 years ago (Newcomer, 1911). Unfortunately, for identification
of the species, he had access only to the color plates of West
Coast butterflies issued by Wright (1907) after the plates of his
book on the same subject had been destroyed in the San
Francisco earthquake and fire of 1906. And since Wright figured
nivalis and called it zeroe, this writer described the early stages
as those of zeroe. Adults were collected at the time and were
later given to the California Academy of Sciences in San
Francisco. Dr. C. Don MacNeill, Assistant Curator, has very
kindly looked these up and he writes about them as follows:
“I find a short series of your specimens of both L. nivalis and
L. mariposa from the Lake Tahoe region. Of your material of
L. nivalis one male and one female each bear an additional hand-
penned label stating zeroe.’ .... All of your material of L.
mariposa came from 5-Lakes^ and none of the specimens bear a
zeroe’ label.”
Later writers, not realizing the misidentification, have con-
sistently said that the early stages of nivalis were unknown.
^These lakes are at an elevation of 7500 feet, tvhile Deerpark, where
nivalis was taken, is at 6500 feet. Both are just south of the Squaw Valley
ski area.
2 (4): 27 1-280, 1963
LYCAENA NIVALIS
^277
Brown (1957) has indicated the range of the subspecies
browni to be southeastern Wyoming, northwestern Colorado and
across southern Idaho; and in a later paper (1961) he has ex-
tended this range to include Washington. In the original descrip-
tion, DosPassos (1938) mentioned Wyoming, Utah, Idaho and
the Methow Valley in Washington. Typical nivalis occurs in
California and up into Oregon.
The subspecies browni occurring in Yakima County, Wash-
ington, dijSers from those collected elsewhere in that the upper
side of the females is usually entirely or almost entirely darkened
by a smoky suffusion that obscures the normal yellowish fawn
ground color. (Since the original description says “females are
very melanic on the upper side,” it may be assumed that this
smokiness occurs elsewhere.) Brown, in his 1961 paper, sur-
mised that this might be due to-, greater humidity in this area.
A study of the Weather Bureau's climatological data indicates
that humidity does apparently affect the coloration but not in the
way suggested by Brown. From available weather records and
with some interpolation, the approximate precipitation in inches
at some of the localities where nivalis occurs may be given
( Table 1 ) . Most of the' precipitation in all of these localities
would be in the form of snow and hence, in addition to the
annual, records are given for April to June, as that would
be the time when larvae and pupae would be present and the
precipitation then would come mostly as rain.
TABLE 1
Locality Elevation Precipitation in Inches
April
May
June
Annual
Lakeview, Lake Co, , Or eg.
4800
L 2
1. 5
1. 4
14
Ochoco Mts, , Crook Co. ,
Or eg.
5000
1,2
2.4
1. 5
19
Tombstone Prairie, Linn
Co. , Oreg,
4000
3, 5
4, 3
1. 0
90
Satus Pass, Klickitat Co.,
Wash,
3700
2, 2
1.2
0. 2
26
Bear Canyon, Yakima Co,,
Wash,
3000
1. 0
0, 8
1. 2
19
Methow Valley (Winthrop),
Okanogan Co* , Wash,
3200
0, 7
1.0
1. 3
14
Harts Pass, Okanogan Co,,
Wash,
5500
6, 0
4. 0
3. 0
85
278
NEWCOMER
/. Res. Lcpicl.
The writer found females of nivalis {zeroe) ovipositing on
Polygonum douglasii at Deerpark, near Lake Tahoe, California,
in July 1909. He took them to Palo Alto and the eggs hatched
the following February. Since doiiglasii was not available there,
he tried related plants and succeeded in rearing some of the
larvae on a species of Rumex. The young larvae ate pits in
the leaves of this plant but older ones devoured the leaves
entirely.
Since the Canadian Entomologist is generally available in
libraries, it will not be necessary to repeat the rather long
description of the early stages here. A brief summary should
suffice for those who do not have access to the original descrip-
tions.
Egg — Diameter, 0.85 mm. Depressed spheroid, ornamented
with deep, polygonal pits; color pale bluish.
Larva — Four instars; newly hatched a little over 1 mm. in
length; full-grown, 17 mm. Slug-shaped, brownish yellow when
first hatched; later becoming pale green with a dorsal line of
rose or claret color; a narrow whitish line on each side of it;
cervical shield sunken on full-grown larvae, in the shape of a
double diamond; dorsal and lateral hairs brownish; dorsum
covered with small, white, hairless tubercles. Head dark brown.
Pupa— Length 10.5 - 11 mm. Rather stout, but longer for its
breadth than usual with lycaenid chrysalids. Color at first very
pale yellowish-green with a pinkish dorsal stripe; later pale
straw-yellow with some brown spots. Hairs, resembling min-
iature trumpets, scattered thickly on dorsum of head and more
sparingly on thorax and abdomen, appearing like minute tacks
stuck into the skin.
In July, 1962, the writer saw a female nivalis ovipositing on
a small, more or less dried-up plant near Satus Pass, Washington.
This plant proved to be Polygonum douglasii,^ an annual. The
^Detennined from Abrams “Illustrated Flora of the Pacifie States,” vol. II,
Stanford Univ. Press, Stanford, Calif., 1944.
area in Bear Canyon, west of Yakima, Washington, where nivalis
is particularly abundant, was immediately examined, and plants
of P. douglasii were found. This species occurs from British
Columbia to the mountains of southern California and east to
the Rocky Mountains. It is sometimes the dominant annual in
high, open prairies. Later, it was found growing wherever
nivalis was flying in Washington and Oregon, and it is, as prev-
iously mentioned, probably the major food plant of the species.
2 (4):27 1-280, 1963
LYCAENA NIVALIS
279
The adults of nivalis frequent the flowers of yarrow {Achillea
millefolmm) , woolly-sunflower {Eriophijllinn lanatwn), spiraea,
and occasionally others in the vicinity of the larval food plant.
Females were collected alive and put in plastic ice-cream
cups with the food plant; and eggs were deposited on the plant
and also on the sides of the containers. These eggs were kept
over winter outdoors but they did not hatch. Probably this was
due either to lack of moisture or to too much cold. The winter
temperature got as low as — 3°F. In the natural habitat the
wintering eggs would almost always be covered with snow
during the cold weather and would therefore not be subject to
temperatures much below freezing. Probably the eggs could
be carried through successfully in a refrigerator.
Thus we see that this single-brooded species passes about
eight months of its life in the egg stage which is subjected to
the heat of summer,— and the eggs would often be on or near
dry ground in full sunlight, — the cold of autumn and the “re-
frigeration” of winter. They would also be subjected to con-
siderable extremes of moisture during that time. And when the
Liow has gone, the newly hatched larvae must seek out the
seedling plants sprouting from seeds produced by the plants
on which oviposition occurred. And yet the species is successful
and is quite common at times.
REFERENCES
BARNES, WILLIAM & McDUNNOUGH, J. 1917. Check List of
Lepidoptera, Herald Press, Decatur, III. p. 15.
BOISDUVAL, J. B. A. 1869. Lepidopteres de la Californie. Ann. Sac.
Ent. Belgique, 12:44-45.
BROWN, F. MARTIN. 1957. Colorado Butterflies. Proc. Denver Mus.
Nat. Hist., Denver, p. 157.
1961. A note about Lycaena nivalis browni ( Lycaenidae) .
Journ. Lepid. Soc. 15:107-8.
CLENCH, HARRY K. 1961. Tripe Lycaenini. In Ehrlich & Ehrlich,
How to Know the Butterflies. Brown, Dubuque, Iowa. p. 226.
COMSTOCK, J. A. 1927. Butterflies of California. Los Angeles, p. 174.
DOS PASSOS, CYRIL F. 1938. Some New Subspecies of North American
Lycaenidae. Can. Ent. 70:45.
DYAR, HARRISON G. 1902. A List of North American Lepidoptera.
Bull. 52, U. S. N. M. p. 41.
EDWARDS, W. H. 1871. Trans. Am. Ent. Soc. 3:211.
1877. Catalogue of the Lepidoptera of America North of
Mexico. Part 1, Diurnals. Am. Ent. Soc. p. 43.
HOLLAND, W. J. 1904. The Butterfly Book. Doubleday Page & Co.,
New York. pp. 251-5.
1931. The Butterfly Book (revised). Doubleday & Co., Garden
N. Y. p. 249.
280
NEWCOMER
/. Res. Lepid.
LEIGHTON, BEN V. 1946. The Butterflies of Washington. Univ. Wash.
PubL in Biology. 9:47-63.
LLEWELLYN JONES, J. R. J. 1951. An Annotated Check List of the
Macrolepidoptera of British Columbia. Occ. paper no. 1, The Ent.
Soc. of B. C. p. 20.
NEWCOMER, E. J. 1911. The Life Histories of Two Lycaenid Butter-
flies. Can. Ent. 43:83-88.
WRIGHT, W. G. 1907. Colored Plates of the Butterflies of the West
Coast. San Bernardino, Calif. Plate XXVIII, no, 346.
Journal of Research on the Lepidoptera
2(4) :281-288, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Ccpyright 1964
COMPARISON OF THE SELECTIVE EFFECT
OF TWO MUSTARD OILS AND THEIR
GLUCOSIDES TO PIERIS LARVAE
WILLIAM HOVANITZ AND VINCENT C. S. CHANG
California Arboretum Foundation, Inc., Arcadia,
and California State College at Los Angeles.
The effectiveness of various mustard oils ( isothiocyanates )
on attracting larvae of Pieris rapae has been shown previously
(Hovanitz, Chang and Honch, 1963b, and Hovanitz and Chang,
1963a). In this paper, some data are presented to compare the
effectiveness of two of these mustard oils with their corresponding
glucosides. The combinations of mustard oils and glucosides
being considered are allyl isothiocyanate with sinigrin, and benzyl
isothiocyanate with glucotropaeolin. The naturally known oc-
currence of the substances in plants is indicated in Table 1 of the
before mentioned paper (Hov., Chang and Honch, 1963b).
THE STRAINS AND EXPERIMENTAL SET-UP
The two strains of Pieris rapae used in these experiments
are the same as used in previous experiments. The first, the kale
strain, has been maintained ten or more generations on kale and
the second, the mustard strain, has been maintained for more than
ten generations on mustard.
The experimental set-up for these tests has been the same as
for those tests previously made (Hovanitz and Chang, 1963a).
ATTRACTION OF GLUCOSIDES
The glucoside, glucotropaelum, is known to occur naturally
in a wide variety of plants, some natural food plants of Pieris
rapae and others not.
Previously published results of tests on the mustard oils cor-
responding to these glucosides have indicated that larvae of
Pieris rapae bred on mustard have no more selection for benzyl
isothiocyanate than for distilled water, where the dilutions have
This work was done with the aid of a grant from the National Science Foundation.
281
TABLE 1, Attraction of glucosides of various concentrations to Pieris rapae larvae
282
HOVANITZ -AND CHANG
/. Res. Lepid.
Total
300
320
240
860
200
160
160
520
280
320
600
600
260
140
320
720
None
39 13.00
25 7. 81
18 7.50
82 9. 53
o o on
lT) ^ on Lfi
nj V -H*
LD Os] on o
nj — 1 nj vD
m
m ' CO
o'
nj -H
sO m
in rg 00
69 11.50
44 16.92
13 9. 28
40 12, 50
97 13. 47
Water
24 8. 00
34 10.62
32 13.33
90 10.46
12 6.00
13 8. 12
14 8.75
39 J. 50
19 6.78
40 12. 50
59 9. 83
69 11.50
^ t'- sO m
f.-, m Lo o
fsj* 00 nO O'
rg nj -H in
m -H (NJ o
56 18.67
49 15.31
33 13.75
138 16. 04
32 16.00
37 23. 12
13 8. 12
82 15.76
48 17. 14
47 14.69
95 15.83
"7 no (O' 00
'O 00 -H cn
fNj’ (NJ* -h"
m 00 O' (Nj
r\] ,_H m 00
00
b
49 16. 33
51 15.94
37 15.42
137 15.93
27 13.50
25 15,62
24 15.00
76 14.62
42 15.00
63 19.69
105 17. 50
40 15.38
26 18.57
51 15.94
117 16.25
b
61 20.33
51 15.94
45 18.75
157 18.25
34 17.00
22 13.75
22 13.75
78 15.00
00 in o
nj m
vD vO vD
r- nj O'
m O'
o
o
00
40 15.38
34 24. 28
48 15.00
122 16.94
'sO
b
ro un — 1 LD
on
fM on m on
^ t- -H
^ on
50 25. 00
27 16.87
28 17. 50
105 20. 19
00
00 -H
sO rg
O' ^ m
•— 1 Lf) f'-
98 16.35
44 16.92
23 16.43
44 13.75
111 15.41
m
b
34 11.33
66 20. 63
38 15.83
138 16. 05
O O O 00
o o m rO
o Ln (N3 m
O o o
(M fN3 CO 00
Tf t'- m
-H 00 m
t'.* -H
00 00 vO
^ m 00
102 17.00
35 13.46
14 10.00
77 24. 06
126 17. 50
97 16. 16
cn
b
81 13.50
Test
1
2
3
Total
1
2
3
Total
1
2
Total
3
1
2
3
Total
jglucoside: |
glucotro-
paeolum
larvae:
mustard-
strain
glucoside:
glucotro-
paeolum
larvae:
kale-
strain
glucoside;
sinigrin
larvae:
mustard-
strain
glucoside:
sinigrin
larvae;
kale-
strain
2 (4):281-288, 1963
GLUCOSIDES
283
ranged from 10"^ to 10 ®. On the other hand, similar larvae tested
for allyl isothiocyanate showed a strong selection for this mustard
oil at a concentration of 10"® (Hovanitz, Chang and Honch
1963b). Larvae bred on kale tested in the same way showed a
decided preference for a more dilute concentration, 10 ® (Hov-
anitz and Chang 1963a ) .
Tests of larvae bred on kale, and other tests of larvae bred on
mustard, were made on the two glucosides ( Table 1 ) . These tests
have indicated that both glucosides have same selective influence
on the larvae but that the selection is not very much greater than
that on water alone.
Glucotropaeolum tested with mustard-strain larvae at con-
centrations of 10‘® to 10‘® and distilled water showed an average
selection of 18 percent for a concentration of 10'^ as compared
with water at 10 percent (Fig. 1). The other concentrations were
not much different at 13 to 16 percent. This compares with no
apparent selection for the corresponding mustard oil, benzyl-
isothiocyanate.
Glucotropaeolum, tested again with kale-strain larvae at
the same concentrations, showed the highest rate of selection,
20 percent, at 10'® as compared with water at 7.5 percent and
other concentrations at 15 percent (Fig. 2). This indicates a
positive selection for the glucoside, whereas no selection was
indicated for the corresponding mustard oil, benzyl isothiocya-
nate.
Sinigrin, the glucoside of allyl-isothiocyanate, was tested
with mustard-strain larvae at concentrations of 10“® to 10"® and
distilled water in a series of two tests and at concentrations of
10"® to 10"^ in a third. The average of the first two tests indicate a
high selection of 17.5 percent at a concentration of 10"® as com-
pared with water at ten percent though other concentrations
were not greatly difiFerent, at 12 to 16.5 percent (Fig. 3). It is
possible that there is little significant effect of change of con-
centration.
The same tests were made over to see if a higher concentra-
tration of sinigrin had a greater influence on selection. Con-
centrations were arranged from 10"® to 10"^ (Fig- 5). The highest
selection, in this series, 17 percent, was at 10 ®, with other
concentrations ranging from 13.5 to 16 percent, compared with
water alone at 11.5 percent. There seems to be little difference in
selection at the different concentrations except that water is
selected less than the glucoside sinigrin. These results differ
PERCENT SELECTION PERCENT SELECTION
284
HOVANITZ AND CHANG
/. Res. Lepid.
Fig. 1. The percent selections by larvae of Pieris rqpae of a standard bred
strain tested for attraction to various dilutions of glucotropaeolum.
Fig. 2. Similar selections by larvae from kale bred strains of Pieris rapae
tested to glucotropaeolum.
PERCENT SELECTION PERCENT SELECTION
2 (4);28 1-288, 1963
GLUCOSIDES
285
Fig. 3. The percent selection by larvae of mustard strain Pieris rapae tested
for attraction to various dilutions of sinigrin.
Fig. 4. Similar selections by larvae of kale strain Pieris rapae to sinigrin.
PERCENT
286
HOVANITZ AND CHANG
/. Res. Lepid.
markedly from the selection data of mustard-strain larvae for
allyl-mustard oil, with a high selection of 30 percent at 10"®
(Hovanitz and Chang, 1963b).
Sinigrin was also tested with larvae from the kale strain
(Fig. 4). The concentration of -0"^ led to selections of 17.5 per-
cent followed closely by 17 percent at lO'h This corresponded to
water at 9 percent, and a range of 11 to 16 percent for the other
concentrations. These results also differ markedly from the
selection data of kale strain larvae for allyl-mustard oil, with
a high selection of 34 percent at 10“® (Hovanitz and Chang
1963a).
CONCLUSIONS AND DISCUSSION
The results of these experiments lead to the following
conclusions :
1. Glucotropaeolum is more effective in attracting larvae of
Pieris rapae than its corresponding oil, benzyl-isothiocyanate.
However, the oil has no apparent attractive influence at all
and the glucoside has only a slight effect.
2. There is little difference in selection between the various
concentrations of the glucoside.
Fig. 5. The percent selections by larvae of mustard strain Pieris rapae
tested for attraction to various dilutions of sinigrin.
2 (4):281-288, 1963
GLUCOSIDES
287
3. There is little difference in selections made by the larvae
toward glucotropaeolum depending on whether they were
from the mustard strain or the kale strain, though it is possibly
significant that a higher selection may occur toward a con-
centration of 10"^ for the mustard strain larvae as compared
with a concentration of 10'® for kale strain larvae.
4. Sinigrin is less eflFective in attracting larvae of Pieris rapae
than its corresponding oil, allyl isothiocyanate.
5. There is little difiFerence in selection between the various
concentrations of the glucoside.
6. There is little difference in selections made by the larvae
toward sinigrin depending on whether they were from the
mustard strain or the kale strain. It is possibly significant that
a higher selection may occur for a higer concentration of
sinigrin for the mustard-strain larvae. This result is op-
posed to the selection data obtained for the same strains
tested for allyl mustard oil ( Hovanitz and Chang, 1963a ) .
Interpretation of these results may depend finally upon future
experiments which would differentiate the ability of the larvae
to perceive odors from a distance. The experimental set-up for
these tests has been based upon the ability of the larvae to react
to odors from a distance of over 100 mm. If the attraction of
larvae to the glucoside sinigrin is really based upon attraction
to the allyl mustard oil vapors emanating from sinigrin by hy-
drolysis, then there may be very little difference in the total
amount of the mustard oil present at any time and this could
conceiveably account for the almost lack of selective differentia-
tion between various concentrations of sinigrin, unlike that of
the mustard oil itself. If the mustard oil emanating from the
sinigrin were restricted by the speed of the reaction of hydrolysis,
then the oil vapors would possibly be dissipated quickly in the
air and not carry far enough to attract the larvae differentially.
On the other hand, selections made close at hand, within a few
millimeters, might serve to bring about such a differentiation.
The lack of any clear-cut selective differences under the con-
ditions of these experiments is strange in view of the great effec-
tiveness of larval selections of different food plants from the same
distances. It is possible that the plants have a means to store
the mustard oils obtained by enzymatic hydrolysis of sinigrin in
greater concentrations that is possible where the sinigrin is
288
HOVANITZ AND CHANG
J. Res. Lepid.
directly exposed to the air. It is diflRcult experimentally to control
the concentration of mustard oils for experimental purposes be-
cause of their great volatility.
LITERATURE CITED
HOVANITZ, W. and V. C. S. CHANG, 1963. a. Selection of allyl isothio-
cyanate by larvae of Pieris rapae and the inheritance of this trait.
/. Res. Lepid. 1 (3): 169-182.
HOVANITZ, W., V. C. S. CHANG and G. HONCH, 1963. b. The
effectiveness of different isothiocyanates on attracting larvae of Pieris
rapae. J. Res. Lepid. 1 (4): 249-259.
Journal of Research on the Lepidoptera
2 (4) : 289^300, 1963
1140 W. Orange Grove Ave., Arcadia, California, U.S.A,
© Copyright 1963
THE ANNAPHILA ASTROLOGA COMPLEX
WITH DESCRIPTIONS OF THREE NEW SPECIES
FRANK P. SALA
3830 Carnavon Way, Los Angeles 27, California
The species-complex presently known as Annaphila mtrologa
Barnes and McDunnough has had a long and confused history.
This paper is an attempt to clarify the status of the several en-
tities now bearing the name A. astrologa B. & McD.
The original type specimens to which Barnes and McDun-
nough ( 1918 ) applied the name A. astrologa were three females,
from various non-specific locations in Arizona. When Riiidge and
Smith (1952) revised the whole genus Annaphila, astrologa was
further described, including the picturing of the genitalia of the
adult male and female, based on an extended series. While other
species were treated in great detail, the authors noted that more
specimen data in the species astrologa might alter their opinion.
This author discovered that it was possible to separate out
from this long series of presumed astrologa several segrates all
of which seemingly fit the description of astrologa. Two of these
were collected together in various locations of Los Angeles,
Orange and Riverside Counties of southern California, A third,
unlike either of the two others, but related to those above, was
collected in the Kern River basin of the Sierra Nevada Mountains.
Through the cooperative assistance of Lloyd Martin of the
Los Angeles County Museum, E. A. Todd of the United States
National Museum (USNM), Frederick Rindge of the American
Museum of Natural History, and Jerry Powell of the University of
California, Berkeley (UCB), the bulk of the material used in the
Rindge-Smith study was reassembled and restudied with the
intent of clarifying the status of these forms.
This work now being completed, the writer is led to the
following conclusions: (1) that the original series used in the
description of astrologa by Barnes and McDunnough included
two species, (2) that the revision of Rindge and Smith (1952}
failed to make this distinction, and working on this duahas-one
thesis, they placed two additional species under the name
mtrologa.
289
290
SALA
J. Res. Lepid.
Fig. 1. Upperside facies of Anaphila astrologa, A. vivianae, A. pseudo astrologa, and A. olgae as indicated.
2 (4):289-301, 1963
ANNAPHILA ASTROLOGA
291
The initial task of the author was to establish which of the
four species was to be called astrologa. In the original series of
three specimens, two are con-specific, the third distinct, as noted
above. One of the former had been used by Barnes and McDun-
nough (1918) to illustrate astrologa. In addition, con-specific
material from California was used by Rindge and Smith to illus-
trate the male genitalia. Thus these two originals are designated
the type series of astrologa, with the TYPE specimen so labled
in the USNM collection. This example most closely agrees with
the original description. The example illustrated in this paper
(Fig. 1, 2) is from California, as are the specimens used in the
drawings of the genitalia (Fig. 3, 4).
Annaphila astrologa B. & McD. (modified description)
(fig. 1, 2, 3, 4).
Wing expanse: 17-20 mm. Head - Vertex: Black, few gray and whitish
hair-like scales. Front: Black, with triangular tuft of long hair-like scales.
Antennae: Black, with whitish scales at segment ends. Palpi: Mostly
white. Thorax: Mostly black, few lighter scales (dorsal). Lighter ventrad,
overlain with lighter hair-like scales of legs. Abdomen - dorsal: black,
white rings at intersegmental area; ventral: same, with longer inter-
segmental hair-like scales giving a lighter aspect. Wings - Fore wing upper:
Ground color black, with scattered lighter scales giving frosted aspect to
base and sub-occipital areas. Basal area black, strongly frosted with light
scales from base to basal line. Basal line whitish to basal dash, which is
also- whitish, indistinct. T. A. line white, “W” shaped, accented basally,
distad at radius, and Cu area, mesad in medial and anal margin areas.
Orbicular indistinct, oval, gray on black, often absent. Ciaviform absent.
Median line broad, black, sub-parallel to T. A. line. Reniform outlined
in white, kidney shaped, gray-cream in center. Connected to costa by
white patch width of reniform, distad fusing with anterior end of T. P.
line white, thin, penciled into a script “B” on distad end of Reniform,
broader posteriad to anal margin. More clearly penciled than in pseu-
doastrologa. Area between T. P. line and S. T. line outlined in black,
frosted with white scales. S. T. Line white, incomplete, broadest at apical
end— faintest at Cu-2. Fringes black except at vein ends, which are white.
Hind wing upper: ground color orange ventrally, bordered in black. Basal
area 1/5 black, often overlain with long orange hairs, giving striated ap-
pearance. Discal spot usually present as lunule or spot (black). Outer
2/5 of wing is black band, broadest as rucurved hook in area of m-1.
Fringes mostly white, black at tips of veins, often 1 mm dark band in area
of anal veins. Fore wing lower: basal area to median line orange . Often
a black area bordered by orange basally and distally. Median line black,
well-marked, often spreading. Widest at costa. Reniform a black dash,
often a black lunule. In field of orange, area between t. p. line and s. t.
line a black band, broadest costad. Tip white. Fringes black, white at
vein ends. Hind wing lower: basal area orange to t. a. line, t. a. line black,
more an arc than above, occasionally incomplete. Discal spot black, lunule,
central area orange to outer margin, outer margin as above. Fringes as
above.
Type: Lectotype, female, Arizona (collector unknown), (no
date), the specimen used is the one Barnes and McDunnough
labeled “Type ” and pictured in their publication in Contr.
292
SALA
/. Res. Lepid.
Fig. 2. Underside facies of Anaphila species as indicated.
5 (4):289-301, 1963
ANNAPHILA ASTROLOGA
293
Nat, Hist. Lepidoptera of N. America, VoL 4, No. 2, PL 19, Fig 11,
This is noted, as the B. & McD, series had as one of the three
specimens the type oiAnnaphila olgae, sp. nov., as defined in this
work.
Early stages: Will be discussed in a ^subsequent paper.
Foodplant: Emmenthalia penduliflora (Whispering Bells) Cali-
fornia foodplant.
Type Locality: Reddington, Arizona.
Range: Southern Arizona; southern California, coastal to San
Francisco, Mojave basin, west side. Not known from the
Sierra Nevada range. This author believes that the Cali-
fornia population, at least, is only on the wing in March
and April. The flowering of growth cycle of the foodplant
( dead in the summer months ) precludes normal flight at
the noted month of July.
Remarks: Rindge & Smith note that more material was needed to
clarify exact relationship in this area. The male genitalia is noted
as variable; in fact, it is very constant, the median spine in the
valve serving as a differentiating character between astrologa
and pseudoastmloga. It also appears that Rindge & Smith pictured
the female genitalia of pseudoastrologa for astrologa, the differ-
entiation here being the markedly stronger sclerotization of the
ductus bursae of astrologa, and its larger size and elongated
posterior cross section. With regard to maculation, astrologa is
best separated by its more definitely penciled design, and pres-
ence of a black reniform spot on the underside of the fore wing.
The latter is often present in vimanae, but the larger size and
lighter total aspect of mvianae definitely separate these species;
the conflict is likely to be with pseudoastrologa, with which it
shares locality in California.
Male genitalia: Figure 3. Valves: symmetrical, complex, elongate, narrow,
vAth costal projection, larger anal projection, tip curved, pointed ( no
ventro-medial projection, as in pseudoastrologa), Tegumen: elongate,
tapering posteriad to base of uncus. Uncus: elongate, tapering posteriad,
tip rounded. Phallus: (expanded). Aedeagus: sclerotized base 2 mm long,
group of 12-15 secondary cornuti on ventral surface. Vesica: armed with
several dozen primary cornuti longer than the secondary group, and
separable into larger basal group (and more numerous filliform distal
group ) .
Female genitalia: Figure 4. Segment 8: A transverse ring, most heavily
sclerotized dorsally, less completely ventrad. Ovipositor lobes: sub-
triangular, scattered setae throughout. Ductus bursae: strongly sclerotized,
more so than any of the other species of the complex, 1.5 mm long. Posterior
end twice as wide as anterior end. Corpus bursae: 2 mm long, surface
with numerous sub-equal small spines.
ovipositor lobt
294
SALA
/. Res. Lepid.
Fig. 3. Female genitalia of Anaphila species as indicated.
2 (4);289-301, 1963
ANNAPHILA ASTROLOGA
295
Annaphila olgae sp, nov. (fig. 3, 4),
The third specimen of the original B. & McD. series is unlike
any other Annaphila known to this writer, and is here named
Annaphila olgae sp. nov., in honor of my mother-in-law, Mrs.
Olga Cornibe Fresco, of South Pasadena, California. While the
writer does not generally approve the naming of single speci-
mens, it is felt that this particular case ought not to be left un-
treated. It does not appear to be a malformed individual, nor
other than a representative example of its kind. Further, it is part
of the original series of the basic species examined (Fig. 1, 2).
Male (unknown). Female: Head - Vertex: black, shaped as astrologa;
front: Mack; antennae: missing; palpi: mostly white. Thorax - Dorsal:
mostly rubbed in holotype, few scales left dark. Ventral: light tan, darker
than any of the other species. Abdomen - Dorsal: a dark brown, under
scope some lighter scales at segment boundaries; rubbed somewhat.
Ventral: lighter than dorsum, but again darker in overall aspect than the
others of the group. Wing span: 16.5 mm (as mounted). Wings - Fore
wing upper: overall most like pseudoastrologa, but with ground color
brown where pseudoastrologa is black. Only exceptions noted. Basal area:
much rubbed, but under scope no evidence of light scales as in either
astrologa or vivianae. Reniform: elliptical, rather than lunar, as in the
others. Hind wing upper: most like vivianae, with basal band and basal
area lighter even than vivianae. Brown again replaces black areas of other
species. Discal spot: absent. Distal band: brown, least wide in m-cu area.
Fringes: brown, white at vein ends. Genitalia: not known; type specimen
not dissected.
Type: Fig. 1, 2. Holotype, female, Arizona (collector unknown) (not date).
The third of the three specimens in the type of series of astrologa, as de-
fined by Barnes and McDunnough. Early stages: unknown. Foodplant:
unknown, but probably an annual in the family Hydrophyllaceae. Type
locality: southern Arizona. Range: unknown. Remarks: this species is
now known by the type alone. Since the specimen is one loaned by the
USNM for this work, it has been returned. Kodachrome slides, with index
to actual size, are available at the Los Angeles County Museum. It is
believed that as the Tucson-Globe area is investigated with respect to the
genus Annaphila, olgae may be taken in this area,
Annaphila pseudoastrologa sp. nov. (fig. 1, 2, 3, 4).
A third species, not part of the original series but included by
Rindge and Smith, is here designated Annaphila pseudoastrologa
sp. nov. Its California range is much the same as astrologa and its
appearance very closely resembles that of astrologa B. & McD. It
is of interest that in the areas where both occur together, examina-
tion of the foodplants growing in intimate contact may often
yield the separate species, each from its chosen plant. The author
has worked out the life cycles of both astrologa and pseudo-
astrologa, but these will be treated in detail in a subsequent paper.
Annaphila pseudoastrologa ii. sp.
Wing expanse 19-24 mm. Male (& Female alike). Head - Vertex: black.
Under scope few gray and orange-brown scales. Front: black, with
triangular tuft of long, hair-like scales. Antennae: gross aspect black, with
medial end of each segment black, distal and white, giving ringed appear-
296
SALA
/. Res. Lepid.
Fig. 4. Male genitalia of Anaphila species as indicated.
2 (4):289-301, 1963
ANNAPHILA ASTROLOGA
297
ance. Palpi: mostly white, with scattering of black, giving a light gray
aspect, longer and lighter than asirologa. Thorax - Dorsal: black through-
out. Ventral: white, especially so due to light hair-like scales on femurae.
Abdomen - Dorsal: black cephalad, with white ring scales on caudal
margin, tliese longer and more profuse than on astrologa. Lateral: transition
between dorsum and ventrum. Ventral: caudal margin scales hair-like,
overlying the anterior darker scales of next posterior segment, giving the
whole surface a whitish aspect. Legs - Segments dark mesad, with distal
light ring, scales hair-like. Wings - wing expanse 19-24 mm. Upper
surface fore-wing: ground color gray-black. Basal line: black, extending
from costa to posterior edge of radius. T. A. line: black, delineated by the
grayish-black mesal area to basal line; zigzag distad on negative veins,
mesad on positive veins. Orbicular: outlined in black, round, center often
gray-black, causing whole area to be indistinct. Claviform: indistinct,
often faintly outlined in black. Median Line: Indistinct to absent; always
most prominent at anal margin. Reinform: White, suboval, lightly tinted
gray central portion, always less so than astrologa; attached to costa by
white patch terminating at t.p. line. T.p. line: white, blending to costal
patch anteriad reniform; a light penciled line forming a script “B,” touch-
ing reniform upper, central, and lower distal points; becomes broadened
between cubitus and anal margin. S.t. line: at apex, a light dash; in-
distinct to absent posteriad r-2 vein. Fringes: black, often lighter at vein
ends. Upper suriace hind icing: General pattern: 1/3 area centrally
orange, border black, costal, distal and anal margins more pronounced than
basal margin. Basal area: scales hairlike, mostly black. T.a. line: in-
distinct, always the margin of bordering band, blending to ochrous center;
always thinnest near end of a-1 vein. Discal spot: usually absent; when
present a dot, rarely a fain dash. Discal-anal border: black, with outline
of mesal edge distinctly less recurved than in astrologa; usually less wide
than in astrologa. Fringes: mostly dark, white at vein ends; occasionally
light anteriad m-1. Lower Surface fore wing: ground color orange; orange
to medial line. Median line: black; variable as to width and degree of
spreading mesad and distad into orange field; question mark-shaped, com-
plete from costa to anal margin; some specimens show costal black dash
marginal scales to median line. Reniform absent. T.p. line: black band,
curved distad from costa to anal margin, broadest at costa: fused to distal
margin between m-2 to cu-2 veins. Tip: white, edged in black. Fringes:
black, white at vein ends. Lower surface hind icing: ground color con-
colorous ochre and black with fore wing. T. a. line: black, variable from
complete to discontinuous line, often variably fused mesad into basal area.
Discal spot: faint to absent in lighter specimens; a few black scales when
present; always less than in astrologa and fainter than in vivianae. Marginal
band: typical for all the complex; black, similar to astrologa, but never
penciled as sharply as in astrologa. Tip: white-yellow twin patch of scales,
edged in black. Hind wing lower surface: tip - fringes black and white,
variable, mostly white near apex. Holotype: Male (Fig. 1) above Seminole
Hot Springs, Coll. F. P. Sala, Santa Monica Mts., L. A. Co., Calif. March
9, 1957. Allotype: Female (Fig, 1) above Seminole Hot Springs, Coll.
F. P. Sala, Santa Monica Mts., L. A. County, Calif. March 10, 1958.
Early Stage: Will be discussed in a subsequent paper. Food plant: Phacelia
parryi Torr. and P. minor Harvy. Type locality: upper Latego Canyon,
Santa Monica Mountains, Los Angeles ( above Seminole Hot Springs )
County, Calif. Range: southern California. The Santa Monica coastal
mountain ranges. The intennediate ranges of the San Gabriels, the San
Bernardino and Santa Rosa ranges, and into tlie desert ranges, wherever
the annual foodplants. Phacelia parryi Torr. and Phacelia minor Harvy.
The known northern limit thus far is southern Ventura County on the
Pacific Coast, and the San Gabriels on the western margin of the Mojave.
This is subject to change with further investigation. It does not appear to
298
SALA
/. Res. Lepid.
fly with vivianae in tlie southern edge of the Sierras, however. Early Stages:
Life cycle will be detailed in a subsequent publication. On wing from late
March through April, depending on location and altitude, and yearly climatic
situation. Remarks: The holotype and allotype of pseudoastrologa have been
deposited in the Los Angeles County Museum along with several paratypes.
A pair of paratypes have also been placed in the collections of tlie USNM,
Yale University, Cornell University, UCB, and the Neidhoefer collection, and
the collection of William Bauer. A. pseudoastrologa is distinguished from
all other species of this group by: Gross aspect: lack of white sealing on
basal upper fore wing. Lack of median spot on hind wing upper and/or
lower side; generally larger ( 20-23 mm wing span ) . Genitalia ( male ) :
median spine in ventromedial spine near apex of valve. Genitalia (female):
incomplete segment 8, ductus iDursae not so sclerotized as astrologa. Gen-
italia: male, Fig. 3, Valves: similar to astrologa, but with prominent ven-
tromedial spine (fig. 4 “spine”) near apex of valve. Tegumen: elongate,
tapering, constricted anteriad uncus, broadened at tip. Uncus: broad at
base, constricted to 1/2 basal width through posterior three quarters of
length; tip pointed. Phalus (expanded): aedeagus similar to astrologa,
secondary spines (cornuti) larger and fewer in number (6-8). Vesica:
primary spines ( cornuti ) sub-equal tliroughout; long, fillifomi. Genitalia
(female, fig. 4): Segment 8 incomplete ring, less sclerotized throughout
than in astrologa. Ovipositor lobes triangular, rounded at corners, half as
wide at base as length. Ductus bursae well selerotized, slightly tapering
from maximum width at posterior opening, less than 1.5 mm long; sclero-
tization less than in astrologa. Corpus bursae gross aspect much like
astrologa.
Annaphila vivianae sp. nov. (fig. 1, 2, 3, 4).
A fourth species from the Kern River basin of Kern County,
California is yet again unlike any of those previously discussed,
being the most spectacular of the group. This insect is here named
Annaphila vivianae sp. nov., in honor of my wife, Mrs. Vivian P.
Sala. Thus far, the known range of vivianae is restricted to Kern
Canyon area of the Kern River. None of the other species of the
astrologa complex has yet been taken from the southern tip of
the Sierra foothill area. The life cycle of vivianae is also complete,
and will be discussed later with the others.
Annaphila vivianae (original description)
Wing expanse 19-24 mm. Male and female alike. Head - Vertex: black,
witli liberal sprinkling of light scales. Front: black and white. Antennae:
gross aspect black, medial end of segments ringed with white, under scope.
Thorax - Dorsal: black and white, giving a dusty gray aspect gross. Ventral:
white, more so tlian for any other members of this group. Abdomen - Dorsal:
black, with white fringe covering inter-segmental area. Ventral: same,
with longer fringe scales giving gross aspect a whiter tone. Legs: black-
white banded. Wings - Upper surface forewing: ground color mostly white,
black band across the orbicular area. White in basal area due to gross suf-
fusion of basal area to T. A. line, and extensive widening of reniform and
T. P. line. Basal line: white, extending from costa to medium. T. A. line:
white, approximately .5 mm. wide, penciled finely in black mesad, extending
from costa to anal margin. Basal area: white. Orbicular: indistinct to
absent; when present outlined in light scales, Claviform: absent. Reniform:
white, lunar, often without brown-gray lunule in center; strongly fused
2 (4):289-301, 1963
ANNAPHILA ASTROLOGA
299
with white at costa. T. P. line: fused to reniform (white) at costa, “B”
marking finely penciled, fused to reniform lower distal margin, fused out to
S. T. line to anal margin. S. T. linei white, finely penciled to cu 1 fused
to anal margin. Area between T. P. line and S. T. line: black. Apex: black,
strongly overlain with white, frosted. Fringes: black, more strongly out-
lined in white at vein ends. Hind wing upper surface: pale orange, lighter
than any other of the group; banded distally in black, mesally brown-black.
Band broken at distal end basal margin. Orange area covering % surface
(more than any of other species). Basal area: basal band narrow, weak,
long hairlike scales. T. A. line: faint, marks border of orange-yellow
certer. Discal spot: usually absent, very faint dash on strongly marked
specimen. Distal band: black, least wide in area from m-2 to cu-2. Fringes:
white, occasionally few dark scales between veins, (lighter than any other
species of the group). Fore wing lower surface: ground color orange. Costa:
orange. Median line: represented by dark spot on costal margin, quadrate
black spot in area of radial to medial veins. Area between T. P. line and
S. T. line a black band from costa to cu-2. Tip white, veins delineated
with darker scales. Fringes dark, white at vein ends. Hind wing lower
surface: ground color orange. Discal spot faint, black. Outer margin band
incomplete, starting at costal tip, broken at r-1 area, again at m-1 area to
cu-1 continuing to inner margin. Apical tip white, veins darker, fringes
white, darker at vein ends.
Male Genitalia: (Fig. 3) Valves: as per astrologa. Gnathos: as per pseu-
doastrologa (restricted caudad). Uncus: as per pseudoastrologa. Phallus
(expanded): aedeagus sclerotized base 2 mm. Ig. group of 6 to 8 secondary
cornuti. Vesica: armed with several dozen primary cornuti subequal and
strongly sclerotized. Female Genitalia (Fig. 4): Segment 8: incomplete
ring. Ovipositor lobes: trangular, rounded, most nearly like astrologa.
Ductus bursae: well sclerotized throughout, less than astrologa approx.
1 mm long, 0.6 mm. wide; less taper than either astrologa or pseudo-
astrologa. Corpus burae: like astrologa and pseudoastrologa.
Type holotype male: (Fig. 1, 2) Lower Kem Canyon, Kern County, Cali-
fornia, March 12, 1956. Coll. Vivian Sala; deposited in Los Angeles County
Museum collection. Alotype female: (Fig. 1, 2): Lower Kern Canyon, Kem
County, California. March 2, 1960. Coll. F. P. Sala; deposited in Los
Angeles County Museum collection. Early stages: Will be discussed in
subsequent paper. Foodplant: P. cicutaria. Type locality: Kern Canyon,
Kem County, California. Range: Southern tip of Sierra Nevada Mountains-
Kern River drainage area canyons. The range may be extended to the
north on the western slope of the Sierra, wherever P. cicutaria occurs in
favored localities. Flight is restricted to a 3-4 week period usually be-
ginning in the middle of March and completed by the second week in
April, varying with the termination of winter rains.
Remarks'.A. vivianae is one of the spectacular species. It is dis-
tinguished from all other Annaphila by the whitish suffusion over
the normally black areas, and in the predominance of orange on
the undersides of the wings. The known range is restricted (thus
far) to the southern Sierra mountain range and vivianae does not
with the other species of the astrologa group. In the Rindge-Smith
material was -one specimen probably from the Havilah, Kern
County, California area, probably a Henry Edwards specimen.
A. vivianae may be the originating stock species. In habit and
early stages, viviane is most like pseudoastrologa. With respect
to sequencing of appearance of the Annaphila species of the Kern
300
SALA
/. Res. Lepid.
Canyon area, it is an earlier flier than are either astrologa or
pseudoastrologa are in their areas.
This species is named for my wonderful wife, Mrs. Vivian
Fresco Sala, who collected the holotype.
Evaluation of the several separating characters of the species
lead me to believe that A. vivtanae might be the forerunner of the
group, pseudoastrologa and olgae being the most closely related,
and astrologa possibly the latest derivative.
A. pseudoastrologa has the widest range in southern Cali-
fornia following the range of its foodplants ( Phacelia parryi Torr.
and F. minor Han^y ) . It has been taken along the Pacific Coastal
plain as high as 5,000 ft. in the coastal and intermediate ranges,
and occasionally in the desert mountains of southern California
and the Arizona types. A. olgae is known only from the type, and
its data is only “So. Arizona, poling” (no date). It does carry a
Poling label, and thus might be from the Tucson area of Arizona.
The rearing of the three species noted has thus far indicated
a marked constancy of maculation and gross structure for each
species. Thus, differences not in accord with these, and from
ecologically separable areas as well, are suspect. Several speci-
mens in the series evaluated as astrologa by Rindge and Smith
from this southern California area do not fit any of the four
species discussed in this paper. These have been separated from
the astrologa series, and while the Rindge-Smith label has been
left on the pin, it has been folded and noted as not agreeing with
the descriptions listed. Two specimens so noted are from “western
San Diego County, California.” They are quite worn and indefi-
nite as to collected location data. It was deemed inadvisable to
use these as the basis for description of species. These will be
held pending acquisition of additional material. One specimen
from eastern San Diego is again different, being markedly more
extensive in areas of black intruding on the orange areas. It most
closely resembles pseudoastrologa and possibly is a race thereof.
It has been set aside for future determination.
It is appropriate to discuss the California habitat of these
species and some of the adaptations of Annaphila in general to
this type of environment:
All Annaphila thus far reared by this writer ( 8 species ) have
indicated an annual life cycle and a pupal diapause, with no lay-
over in the pupal stage beyond one year. Thus, in general, a
species can be assumed to be restricted to those areas where the
foodplant, an annual, is able to grow from seed each year.
2 (4). -289-301, 1963
ANNAPHILA ASTROLOGA
301
It is the nature, in general, for the annual foodplants to be
restricted to canyon slope areas, and to grow most profusely
where the ground is often naturally disturbed (silt-flooding or
slope slippage). These plants are clustered in chosen locations
and found not at all elsewhere.
The various California mountain ranges, within which these
foodplant locations are found, are strongly separated one from
another ecologically. Add to this the local nature of the flight
patterns of Annaphila, and the probability of transfer of the
adults from one location to another is small indeed.
Degrees of isolation can be noted. The Sierra flora and ac-
companying Annaphila are strongly divergent from all the other
coastal and intermediate southern California ranges. A. vivianae
is not known to occur with any other of the astrologa complex
species.
Further, the several coastal and intermediate ranges are now
ecological islands as well. A. astrologa and pseudoastrologa fly
together in the Santa Monica and the San Gabriel Mountain
ranges, and thus these two are sympatric.
Acknowledgments
In addition to the aforementioned assistance in loaning of the
materials used in this work, the assistance of Dr. Leland R.
Brown, of the Department of Entomology of the University of
California, and the Department of Entomology of the Los Angeles
County Museum is gratefully noted.
Finally, the author is most grateful to Dr. Charles L. Hogue of
the Los Angeles County Museum, who unselfishly assisted in the
preparation of the illustrations and suggested the method for
preparation of the genitalia.
Literature Cited
BARNES, W. and J. McDUNNOUGH. 1918. Original Description of
Annaphila astrologa. Cont. Nat. Hist. Lepid. No. America, 4(2):
JEPSON, W. L. 1925. Manual of Flowering Plants. Galifornia School Book
Depository.
RINDGE, F. H. and C. 1. SMITFI. 1952. A Review of the Genus Annaphila
Grote ( Lepidoptera, Philadelphia). Bull. Amer. Mus. Nat. Hist.
98(3): 191-256.
302
}. Res. Lepid.
OLD MASTER PAINTINGS
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Eleven painted panels, 48 inches by 65 inches overall.
To be auctioned. For information contact CLARK, NELSON of LONDON LTD., 717
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mailed at the post office at Sierra Madre, Calif.
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Journal of Research on the Lepidoptera ■ 2 (4) : 303-304, 1963
1140 W. Orange Grove Ave., Arcadia^ California, U.S.A.
© Copyright 1963
COLLECTING OF ANNAPHILA SPILA
WITH NOTES ON THE “CRIMSON-WINGED” GROUP
OF THE GENUS
J. S.- BUGKETT
University of California,
Davis, California
The largest series to date of Annaphila spila Rindge &
Smith was collected in the spring of 1961, at which time a series
of over forty specimens was taken in a sheltered draw and along
a creek just below Monticello Dam in Solano County, California.
In this area, erogeny is magnificently pronounced, with some of
the strata almost perpendicular, and as a result, the hillside
habitat of spila proved very difficult collecting. The area is
predominantly chaperral with a scattering of Pinus sahiniana and
various species of Quercus dotting the hillsides.
The flight time is relative to the morning temperature and
seemed to begin around 10:00 A.M.; no specimens of spila were
collected or seen after 4:00 P.M., even though the temperature
still seemed favorable for their flight. Moderate breezes were
blowing, and in combination with the temperature may have had
some bearing on the termination of the diurnal flight. However,
the breezes were also evident around 12:30 P.M., at the peak of
flight. The reasons for the terminations of the diurnal flight of
spila as well as other species of Annaphila still remains unknown.
Specimens of A. decia Grote, A. depicta Grote, and A. divinula
Grote were collected in the same area until 5:00 P.M., and prob-
ably could have been collected later in the day as well.
The adults of spila were found to frequent the flowers of
a small number of plant species, predominantly Monar della
brew-eri. The adults were also collected on Gilia sp., Erodium
sp., and Fsycocarpos sp. Occasionally they were seen hovering
around Germs occidentalism but none were observed either alight-
ing on the leaves or feeding on the flowers. One specimen was
collected around 12:30 P.M., while apparently drinking water
from a damp, sandy spot along a stream.
The 'Trims on- winged” group in the genus Annaphila is com-
prised of superha Henery Edwards, spila Rindge & Smith, and
evansi Rindge & Smith. A. superha may be readily distinguished
from both spila and evansi by the reniform being centrally of
ground color, outlined in white, and. connected to the costa by a
white patch above the anterodistal margin of the reniform; by the
303
304
BUCKETT
/. Res. Lepid.
ventral surface of the primaries possessing a complete median
cross line or band. A. spila and evansi are more closely related
to each other than either is to superba, as is readily seen by the
maculation of the dorsal surfaces of the primaries; the transverse
ridge across the top of the frons; the female and male genitalia.
Neither spila nor evansi display a white outlined reniform; how-
ever in freshly emerged specimens a faint gray outlining of the
reniform may be seen; nor do they display complete median
cross lines on the ventral surface of the primaries as does superba.
A. spila has a dot on the ventral surface of the primaries in place
of the band present in superba, and evansi lacks both of these
characters.
As may be seen by the relationship in the genitalia of these
species, it seems plausable to assume they arose from a common
ancestor, and that superba is evolving as a discrete unit while
spila and evansi are evolving as another more closely associated
unit. The ‘"crimson-winged” group is most probably an offshoot
of the “astrologa-ida" group. The female genitalia of superba
show a similarity to the female genitalia of ida Rindge & Smith,
but the genitalia of the males do not show this as well.
In the “Revision of the genus Annaphila' (Rindge & Smith,
1952), the new species spila was based on very few specimens of
which only one was a male. A female was chosen as the type
and the description was drawn from this sex. I was not sure of
the justification of including the allotype male in the type series,
as it was collected in an entirely different geographical area
which is separated from the type locality by the great interior
valley of California. Further study has proved the populations
from the coast ranges to be conspecific with those of the foot-
hills of the Sierra-Nevada, just as Rindge and Smith suspected.
In the original description of spila, it was stated that this is
an obscurely marked species. I find this not to be so in every
instance; but it must be kept in mind that Rindge & Smith had
very few specimens. The species of Annaphila are very easily
rubbed, and this probably accounts for spila being described as
an obscurely marked species. However, in comparison with
superba, both spila and evansi in excellent condition are more
obscurely marked; but in fresh specimens one can readily see the
cross lines of the primaries which are light gray to white in
both sexes.
LITERATURE CITED
RINDGE, F. H. and SMITH, C. I. 1952. A revision of tlie genus Annaphila
Grote ( Lepidoptera, Phalaenidae ) . Bull., Amer. Mus. Nat. Hist. 98(3):
57 pp., 8 plates.
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om THE LEPIJDOFTE^A
Volume 2
Number 4
December, 1963
IN THIS ISSUE
Evidence for lack of territoriality in two species of Hamadryas
Gary N. Ross 241
A synopsis of the west Indian Lycaenidae, with remarks on
their zoogeography Harry K. Clench 247
The synonymy, variability and biology of Lycaena nivalis
E. J. Newcomer 271
Comparison of the selective effect of two mustard oils and
their glucosides to Pieris larvae
W. Hovanitz and V.C.S. Chang 281
The Annaphila astrologa complex, with descriptions of three
new species Frank P. Sala 289
Old master painting Clark, Nelson of London, Ltd. 302
Dates of publication of volumes 1 and 2 302
Collecting of Annaphila spila with notes on the "crimson-
winged” group of the genus J. S. Buckett 30-3
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